Full text of "Apex"
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HARVARD UNIVERSITY
e
Library of the
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MOL/IS: (1-2)
B. Tursch
D. Greifeneder
D. Huart
J.-M. Pacaud
R.A. Van Belle
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E. Guillot
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SOMMAIRE
A puzzle of highly multiform species :
Oliva fulgurator (Rôding, 1798) and related American taxa.
Un nouveau genre de Pholadidae (Mollusca, Bivalvia)
du Paléogène inférieur du nord de l'Atlantique
Description of a new species Zschnochiton dolii sp. nov.
(Polyplacophora: Ischnochitonidae) from Civitavecchia, Italy
Description d’une nouvelle espèce de Costellariidae
des Philippines
APEX
Société Belge de Malacologie a.s.b.l.
Editeur responsable:
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Comité d’édition:
Dr. T. Backeljau Koninklijk Belgisch Instituut voor Natuurwetenschappen
Dr. Y. Finet Muséum d'Histoire Naturelle, Genève
M. R Houart Institut royal des Sciences naturelles de Belgique (collab. scient.)
Dr. CI. Massin Institut royal des Sciences naturelles de Belgique
Prof B. Tursch Université Libre de Bruxelles
Dr. J. Van Goethem Koninklijk Instituut voor Natuurwetenschappen
Prof. G. Vauquelin Vrije Universiteit Brussel
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Studies on Olividae
28
À puzzle of highly multiform species:
Oliva fulgurator Rôding, 1798)
and related American taxa
Bernard TURSCH
Dietmar GREIFENEDER
Dominique HUART
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
A puzzle of highly multiform species:
Oliva fulgurator (Rôding, 1798) and related American taxa !
Bernard TURSCH, Dietmar GREIFENEDER * and Dominique HUART.
Laboratoire de Bio-Ecologie, Faculté des Sciences, Université Libre de Bruxelles,
50 av. F.D. Roosevelt, 1050 Brussels, Belgium.
!. This is paper n° 28 in the series Studies on Olividae.
2 Research associate.
KEYWORDS. Mollusca, Gastropoda, Oliva, taxonomy, Western Atlantic, Eastern Pacific,
Caribbean, Panamic.
ABSTRACT. The taxonomic status of the members of the Western Atlantic "Oliva fulgurator-
reticularis complex", the Eastern Pacific "Oliva spicata complex" and of some related taxa 1s
reviewed. ©. foxi Stingley, 1984, ©. fulgurator (Rôding, 1798), ©. polpasta Duclos, 1833, O.
scripta Lamarck, 1811 and ©. spicata (Rôding, 1798) are shown to be distinct species. ©. spicata
deynzerae Petuch & Sargent, 1986 is a distinct subspecies. The available type material of many
synonymous taxa has been studied and illustrated.
RÉSUMÉ. Le statut taxonomique des membres du "complexe Oliva fulgurator-reticularis" de
l'Atlantique-Ouest, du "complexe Oliva spicata" du Pacifique-Est et de quelques espèces
apparentées est revu. ©. foxi Stingley, 1984, ©. fulgurator (Rôding, 1798), ©. polpasta Duclos,
1833, ©. scripta Lamarck, 1811 and ©. spicata (Rôding, 1798) sont démontrées être des espèces
distinctes. ©. spicata deynzerae Petuch & Sargent, 1986 est une sous-espèce distincte. Le matériel
APEx 13(1-2): 1-61, 20 avr. 1998
type accessible de nombreux taxa synonymes a été étudié et illustré.
1. INTRODUCTION
1.1. The problem
1.1.1. The number of species
Authors differ widely in opinion on the number of
Oliva species in the Western Atlantic. ZEIGLER &
PORRECA (1969) as well as ABBOTT (1974) admit the
existence of four species [Oliva fulgurator (Rôding.
1798), ©. reticularis Lamarck, 1811, ©. sayana
Ravenel, 1834 and ©. scripta, Lamarck, 1811].
PETUCH & SARGENT (1986) recognise no less than
twenty-one species and ten subspecies.
Many Western Atlantic taxa share common
characteristics and appear to be closely related, forming
the “O. reticularis complex” of TURSCH & HUART
(1990) (this shall here be called the “O. fulgurator-
reticularis complex”, for reasons that will become
apparent in $ 5.1). The numerous, controversial taxa
erected for the members of this group are the elements
of an old and intricate nomenclatural puzzle, possibly
the best example of the notorious taxonomic chaos
prevailing in the genus Oliva. With the exception of the
new species described by PETUCH & SARGENT (1986)
and by PETUCH (1987,1988,1990), not hitherto revised,
nearly every Caribbean taxon of the complex has
already been synonymised (this will be seen in the
section Systematics). Only ©. fulgurator and ©.
reticularis were generally considered to be distinct
species. The existence, the nature and the difficulty of
the problem raised by the “O. fulgurator-reticularis
complex” were already clearly grasped well over a
century ago:
“Oliva reticularis is, without question, the most
variable species in the genus and we warn
conchologists that one should have access to a
very rich, very abundant collection in order to
understand the unions [of taxa] that we are
suggesting ... Let us repeat that such a work
requires a considerable collection as well as
extreme tenacity".
DUCROS DE SAINT GERMAIN (1857: 53)
(translation ours)
The subsequent naming of many vague and i1ll-
defined forms only added to the confusion. The
difficulties encountered in matching shells with names
are familiar to every student of Caribbean Oliva.
In the Eastern Pacific, ZEIGLER and PORRECA
(1969) recognise nine species, while PETUCH &
SARGENT (1986) list twenty-one species (and six
subspecies). The Eastern Pacific Oliva fauna is much
richer in species than its Western Atlantic counterpart.
It contains a few very distinct species [O. porphyria
(L., 1758), ©. peruviana Lamarck, 1811, ©
splendidula Sowerby, 1825, ©. incrassata (Lightfoot in
Solander, 1786), O. julieta Duclos, 1840, ©. kaleontina
Duclos, 1835, ©. undatella Lamarck, 1811, etc.) which
are outstanding in aspect and pose no serious
identification problem.
In addition, there is a large group of obviously
cognate nominal taxa [the “O. spicata complex” based
1
APEX 13(1-2): 1-61, 20 avr. 1998
upon ©. spicata (Rôding, 1798), see TURSCH & HUART
1990]. This poses much the same problems as its
Western Atlantic counterpart, the “O. /ulgurator-
reticularis complex”.
1.1.2. The distribution of species
The geographical distribution of many named taxa in
both the “O. fulgurator-reticularis complex” and the
“O. spicata complex” is poorly known. The limits of
their variation are indeed so imprecise that correct
identification is often possible only for material from
type localities.
Further taxonomic complications arise because
several cognate O/iva taxa of the two faunas (especially
in the “O. spicata complex” and in the “O. fulgurator-
reticularis complex”) are quite similar in appearance.
Early locality data are often questionable, and in
several instances (see examples in Table 1) authors
have considered a same taxon to have an Atlantic or a
Pacific distribution.
Furthermore, some taxa such as ©. olorinella
Duclos, 1835 (q.v.) have been erected upon a mixture
of Atlantic and Pacific shells.
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
1.1.3. Previous quantitative results.
Using morphometric analysis, TURSCH & HUART
(1990) demonstrated that the Atlantic Oliva specimens
fall into four objective groups. One of these groups is
the West African ©. flammulata Lamarck, 1811. The
three others live in the Western Atlantic: ©. scripta
Lamarck, 1811, ©. sayana Ravenel, 1834 and a highly
variable group then called the "©. reticularis complex"
(now the "O. fulgurator-reticularis complex"). None of
these groups could be split into smaller, objective units,
although the complex contains very dissimilar
populations. The only possible candidate for
segregation was ©. oblonga Marrat, 1870, of which
only a small sample was available to us at the time, so
that no definite conclusion could be drawn.
Most Eastern Pacific specimens were easily
separated into the objective groups ©. porphyria, O.
splendidula, ©. kaleontina, ©. undatella, ©. polpasta
Duclos,1833 and a large, highly variable group called
the "O. spicata complex". Further analysis tentatively
separated only a sample of ©. polpasta from a small
allopatric sample of ©. kerstitchi da Motta, 1985. The
specific status of the latter was considered to be only a
working hypothesis, awaiting further confirmation.
taxon Eastern Pacific distribution Atlantic distribution
for for
©. brunnea Marrat, 1870
O. graphica Marrat, 1870
©. oblonga Marrat, 1870
©. oriola ; Duclos
(not Lamarck,
1835.
1811),
©. oniska Duclos, 1845
©. porcea Marrat, 1870
BURCH & BURCH (1960),
WAGNER & ABBOTT (1978)
(as a synonym of ©. spicata)
BURCH & BURCH (1960)
(as a synonym of ©. spicata)
WAGNER & ABBOTT(1978)
(as a synonym of ©. scripta)
BURCH & BURCH (1960), KEEN
(1971), WAGNER & ABBOTT
(1978)
(as a synonym of ©. spicata)
BURCH & BURCH (1960),
ZEIGLER & PORRECA (1969),
KEEN (1971)
(as a synonym of ©. spicata)
WAGNER & ABBOTT (1978)
(as a synonym of ©. fulgurator)
PETUCH & SARGENT (1986)
(as a subspecies of ©.
tisiphona)
WAGNER & ABBOTT(1978)
(as a synonym of ©. scripta)
Table 1. Some examples of divergent geographical attributions.
PETUCH & SARGENT (1986)
(as a subspecies of ©. jamaicensis)
ZEIGLER & PORRECA (1969),
KEEN (1971)
(as a synonym of ©. julieta) PETUCH
& SARGENT (1986)
(as a full species)
PETUCH & SARGENT (1986),
PETUCH (1987)
(as a form of ©. tisiphona)
WAGNER & ABBOTT (1978)
(as a synonym of ©. reticularis, pars)
ZEIGLER & PORRECA (1969)
(as a form of ©. spicata)
BURCH & BURCH (1960)
(as a synonym of ©. spicata)
ZEIGLER & PORRECA (1969),
KEEN (1971)
(as a synonym of ©.
julieta)
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
It was stressed that nearly all the many controversial
taxa erected for American species appear:
1/ to have a restricted geographical distribution.
2/ to belong either to the "©. fulgurator-reticularis
complex" or to the "O. spicata complex" (see
TURSCH & HUART 1990).
1.2. Aim.
The present paper is the continuation and the extension
of the exploratory quantitative survey of living
American Oliva species by TURSCH & HUART (1990).
In that preliminary study, some species were not
represented, other required larger samples and
nomenclatural problems were not treated in any detail.
The causes of the discrepancy of opinions in the "©.
Julgurator-reticularis complex" and the "O. spicata
complex" will be analysed and discussed. A critical
review of the type material will now be presented.
The very distinct species ©. porphyria, O.
peruviana, ©. splendidula, ©. incrassata, ©. julieta, O.
kaleontina, ©. undatella will be mentioned in the
present paper only as references for the delimitation of
the “O. spicata complex”. The West African species ©.
flammulata will also be considered for comparison
because it is the type species of the genus Strephona
Môrch, 1852, in which many American taxa have been
placed (for reasons not clear to us). The taxa O.
schepmani Weissbord, 1962 [most probably a form of
O. fulgurator (Rôding, 1798)], ©. davisae Durham,
1950 (most probably a form of ©. polpasta Duclos,
1833) and ©. callosa Li, 1930 (a subjective junior
synonym of ©. polpasta Duclos, 1833) all based upon
fossil material, will not be examined here.
2. MATERIAL EXAMINED
2.1. Specimens.
372 specimens, from many localities, have been added
to the 419 specimens measured in detail and listed in
TURSCH & HUART (1990). In addition to the material
measured, several thousand specimens, from a large
number of localities, have been examined in public and
private collections. Broad localities for specimens of
the "O. fulgurator-reticularis complex", the main
object of this study are shown on the map of Fig. 1.
The complete list of the examined material would take
many pages and (contrary to our habit) we have
preferred to list only the type material. 35
measurements (17 used in this paper) have been
determined on each intact specimen measured, so this
work rests upon a databank of well over twenty
thousand high-precision measurements (see section
3.1), accumulated over many years.
re
f
Fig. 1. The “O. fulgurator-reticularis complex”. Localities of material examined. Localities in close vicinity
(less than 20 miles) not separated.
APEX 13(1-2): 1-61, 20 avr. 1998
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
2.2. Abbreviations:
AMS: Australian Museum, Sydney.
AMNH: American Museum of Natural History, New
York.
ANSP: Academy of Natural Sciences, Philadelphia.
BM(NH): The Natural History Museum, London.
MCM: Merseyside County Museum, Liverpool.
MCZ':Museum of Comparative Zoology, Harvard
University.
MNHN: Muséum National d'Histoire Naturelle, Paris.
MANG: Muséum d'Histoire Naturelle, Genève.
USNM: National Museum of Natural History
(Smithsonian Institution), Washington.
SBMNH!: Santa Barbara Museum of Natural History.
SMF: Forschungsinstitut und Naturmuseum
Senckenberg, Frankfurt.
2.3. Type material examined.
Oliva foxi Stingley, 1984.
Oliva foxi Stüngley, 1984. Holotype ANSP 358356 (H:
36.98 mm, D: 14.82 mm) (see PL. 4, FIG. 1). One
paratype ANSP 358357 (H: 37.60 mm, D: 15.17 mm)
(see PL. 4, FIG. 2). One paratype AMNH 264800 (H:
34.9 mm, D: 14.1 mm) (see PL. 4, FIG. 3). One
paratype AMS C.170890 (H: 33.35 mm; D: 13.36
mm) (see PL. 4, FIG. 4).
Oliva fulgurator (Rôding, 1798).
Oliva olorinella Duclos, 1835. Eight heterogeneous
syntypes at MNHN. One syntype (H: 28.7 mm, D:
12.1 mm) illustrated PL. 9, FIG. 2.
Oliva obesina Duclos, 1840. Figured syntype (H: 44.0;
D: 26.3 mm) (see PL. 6, FIG. 1), and 2 syntypes at
MNAN.
O. timoria Duclos, 1840 (pars). Figured syntype (out
of 3) (H: 62.0 mm, D: 33.1 mm), MNHN. (see PL. 7,
FIG. 1).
Oliva aldinia Duclos, 1845. Figured syntype (H: 55.6
mm, D: 30.3 mm), (see PL. 6, FIG. 2) + 3 syntypes at
MNAHN.
Oliva broderipi Ducros de St. Germain, 1857.
Holotype at MNHN (H: 27.8 mm, D: 13.4 mm) (see
PÉwAEG 3):
Oliva jamaicensis Marrat, 1867. Holotype at MCM (H:
37.7 mm, D: 18.4 mm) (see PL. 6, FIG. 6).
Oliva oblonga Marrat, 1867. Holotype at MCM (H:
63.2 mm, D: 29.2 mm) (see PL. 6, FIG. 7).
Oliva bewleyi Marrat, 1870. Holotype at MCM (H:
41.8, D: 18.1 mm) (see PL. 7, FIG. 10).
Oliva figura Marrat, 1870. Holotype at MCM (H: 41.2
mm, D: 18.1 mm) (see PL. 8, FIG. 3).
Oliva formosa Marrat, 1870. Holotype at MCM (H:
45.23 mm; D: 19.52 mm) (see PL. 9, FIG. 6).
Oliva graphica Marrat, 1870. Holotype at MCM (H:
44.6 mm, D: 22.2 mm) (see PL. 6, FIG. 4).
Oliva porcea Marrat, 1870. Holotype at MCM (H: 42
mm, D: 20.8 mm) (see PL. 6, FIG. 3).
Oliva sowerbyi Marrat, 1870. Syntype (out of 2) at
MCM (H:32.2 mm, D:15.0 mm). This taxon is not
related to ©. fulgurator (see Systematics).
Oliva bullata Marrat, 1871. Holotype at MCM (H:
60.83 mm, D: 26.21 mm) (see PL. 9, FIG. 3).
Oliva mercatoria Marrat, 1871. Two syntypes at MCM
(H: 41.0 mm, D: 19.5 mm and H: 44.2 mm, D: 20.5
mm) (see PL. 6, FIG. 9).
Oliva nivosa Marrat, 1871. Holotype (H: 49.2 mm, D:
22.0 mm) (see PL. 7, FIG. 9) and one possible
paratype at MCM.
Oliva reclusa Marrat, 1871. Holotype at MCM (H:
43.2 mm, D: 18.1 mm) (see PL. 9, FIG. 7).
Oliva circinata Marrat, 1871. Holotype at MCM (H:
55.4, D: 23.4 mm) (see PL. 9, FIG. 8), with 4 possible
paratypes. One probable paratype (as stated on an
additional label written by N. McMillan in 1978)
ANSP 15940 (H: 47.02 mm, D: 19.64 mm).
Oliva bifasciata Küster in Weinkauff, 1878. Figured
specimen SMF 9353 (H: 61.39 mm, D: 26.73 mm)
(see PL. 8, FIG. 4).
Oliva cribraria Marrat, 1883? Holotype (H: 48.22 mm;
D: 22.12 mm) at MCM (see PL. 9, FIG. 10).
Oliva reticularis bollingi Clench, 1934. Holotype MCZ
76656 (H: 61.95 mm, D: 28.52 mm) (see PL. 7, FIG.
8).
.Oliva reticularis greenwayae Clench, 1937. Holotype
MCZ 115455 (H: 59.65 mm, D: 25.18 mm) (see PL.
8, FIG. 5).
Oliva pattersoni Clench, 1945. Holotype MCZ 151166
(H: 62.57 mm, D: 28.29 mm) (see PL. 8, FIG. 6).
Oliva drangai Schwengel, 1951. Holotype ANSP
247107 (H: 40.47 mm, D: 18.62 mm) (see PL. 7, FIG.
7). Paratype ANSP 247093 (H: 18.56 mm, D: 8.46
mm).
Oliva antillensis Petuch & Sargent, 1986. Holotype
USNM 841425 (H: 28.05 mm, D: 11.47 mm) (see PL.
9, FIG. 1).
Oliva ionopsis Berry, 1969. Holotype SMBNH 34656
(H: 30.54 mm; D: 13.76 mm) (see PL. 11, FIG. 10)
and seven paratypes SMBNH 34657.
Oliva bahamasensis Petuch & Sargent, 1986. Holotype
USNM 841426 (H: 44.58 mm, D: 20.62 mm) (see PL.
8, FIG. 10).
Oliva barbadensis Petuch & Sargent, 1986. Holotype
USNM 841427 (H: 47.05 mm; D: 20.32 mm) (see PL.
9, FIG. 4).
Oliva bifasciata jenseni Petuch & Sargent, 1986.
Holotype USNM 859302 (changed from USNM
841453, R. GERMON in litt.) (H: 50.30 mm, D: 24.12
mm) (see PL. 7, FIG. 6).
TURSCH, GREIFENEDER & HUART
Oliva finlayi Petuch & Sargent, 1986. Holotype USNM
841432 (H: 39.98 mm, D: 18.71 mm) (see PL. 8, FIG.
8).
Oliva goajira Petuch & Sargent, 1986. .Holotype
USNM 841433 (H: 36.79 mm, D: 15.49 mm) (see PL.
8, FIG. 2).
Oliva jamaicensis zombia Petuch & Sargent, 1986.
Holotype USNM 841454 (H: 23.87 mm, D: 12.61
mm) (see PL. 7, FIG. 2).
Oliva magdae Petuch & Sargent, 1986. Holotype
USNM 841441 (H: 37.57 mm, D: 18.62 mm) (see PL.
6, FIG. 8).
Oliva maya Petuch & Sargent, 1986. Holotype USNM
859301 (changed from USNM 841453, R. GERMON
in litt.) (H: 57.81 mm: D: 24.97 mm) (see PL. 8, FIG.
1).
Oliva bifasciata sunderlandi Petuch, 1987. Holotype
USNM 859904 (H: 22.02 mm, D: 9.56 mm) (see PL.
9, FIG. 5).
Oliva circinata tostesi Petuch, 1987. Holotype USNM
859865 (H: 46.20 mm, D: 21.37 mm) (not H: 41 mm
as in description) (see PL. 9, FIG. 9).
Oliva sargenti Petuch, 1987. Holotype USNM 859864
(H: 28.60 mm, D: 14.03 mm) (see PL. 7, FIG. S).
Oliva contoyensis Petuch, 1988. Holotype USNM
859945 (H: 36.71 mm, D: 18.42 mm) (see PL. 7, FIG.
4).
Oliva ernesti Petuch, 1990. Holotype USNM 860535
(H: 33.66 mm, D: 15.45 mm) (not H: 38 mm, as in
description) (see PL. 8, FIG. 7).
Oliva polpasta Duclos, 1833.
Oliva polpasta Duclos, 1833. Six syntypes at MNHN.
Figured syntype (H: 49.1 mm; D: 21.0 mm) (see PL.
4, FIG. 6).
Oliva truncata Marrat, 1867. Holotype at MCM (H: 37
mm, D: 18 mm) (see PL. 4, FIG. 7).
Oliva kerstitchi da Motta, 1985. Holotype MHNG
984.631 (H: 24.5 mm, D: 11.2 mm) (see PL. 4, FIG.
8).
Oliva olssoni Petuch & Sargent, 1986. Holotype
USNM 841444 (H: 35.03 mm; D: 19.57 mm) (see PL.
4, FIG. 9).
Oliva sayana Ravenel, 1834.
Oliva circinata var. citrina Johnson, 1911. Paratype
MCZ 6267 (H: 60.51 mm, D: 23.56 mm) (see PL.S5,
FIG. 4).
Oliva sayana sarasotensis Petuch & Sargent, 1986.
Holotype USNM 841450 (H: 40.60 mm, D: 17.90
mm) (see PL. 5, FIG. 3).
Oliva sayana texana Petuch & Sargent, 1986. Holotype
USNM 841465 (H: 43.60 mm, D: 17.54 mm) (see PL.
5, FIG. 2).
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
Oliva scripta Lamarck, 1811.
Oliva caribaeensis Dall & Simpson, 1901. Holotype
USNM 159672 (H: 31.61 mm, D: 14.93 mm) (see PL.
5, FIG. 6).
Oliva trujilloi Clench, 1938. Holotype MCZ 57240 (H:
40.52 mm, D: 17.95 mm) (see PL. 5, FIG. 7). One
paratype AMNH 79010 (H: 34.8, D: 15.5 mm) (see
PL. 5, FIG. 8). Five paratypes AMS C.095147 (one H:
39.57 mm, D: 18.59 mm, illustrated PL. 5, FIG. 9).
Oliva (Cariboliva) scripta venezuelana Petuch &
Sargent, 1986. Holotype USNM 841451 (H: 44.22
mm, D: 22.02 mm) (see PL. 5, FIG. 10).
Oliva spicata (Rôding, 1798).
Oliva oniska Duclos, 1845. Nine syntypes at MNHN.
Figured syntype (H: 61.0 mm; D: 26.2 mm) (see PL.
11, FIG. 4).
Oliva pindarina Duclos, 1840. Three syntypes at
MNAN. Figured syntype (H: 53.4 mm; D: 27.3 mm)
(see PL. 10, FIG. 2).
O. timoria Duclos, 1840 (pars). Figured syntype (out
of 3) (H: 55.8 mm, D: 39.1 mm), MNAHN. (see PL.
10, FIG. 1).
Oliva subangulata Philippi, 1848. One syntype (H:
59.62 mm; D: 26.38 mm) out of 2, BM(NH)
1924.1.5.98-99. (see PL. 11, FIG. 6).
Oliva cumingii Reeve, 1850. Three syntypes BM(NH)
1987008, (one H: 48.4 mm, D: 24.2 mm) (see PL. 11,
FIG. 1).
Oliva ligneola Reeve, 1850. Two heterogencous
syntypes BM(NH) 1892.9.24.4-5. One (H: 21.36 mm;
D: 10.67 mm) is ©. tigrina fallax Johnson 1911, the
other (H: 25.60 mm, D: 13.09 mm) (see PL. 10, FIG.
3) is a bleached ©. spicata (Rôding), 1798.
Oliva intertincta Carpenter, 1857. Two syntypes
USNM 716187 (largest: H: 19.68 mm, D: 9.70 mm,
see PL. 11, FIG. 9). 3 additional syntypes in BM(NH),
Carpenter collection, tablet 2121.
Oliva violacea Marrat, 1867. Holotype at MCM (H:
39.3 mm, D: 17.3 mm) (see PL. 11, FIG. 8).
Oliva brunnea Marrat, 1870. Holotype at MCM (H:
34,0 mm, D: 16.1 mm) (see PL. 11, FIG. 3).
Oliva punctata Marrat, 1870. One of two syntypes (H:
35.35 mm, D: 17.24 mm) at MCM (see PL. 10, FIG.
4).
Oliva fuscata Marrat, 1870. Three syntypes at MCM,
one of which (H: 42.66 mm, D: 20.30 mm) is figured
PL. 11, FIG. 2 and another (H: 39.76 mm, D: 18.62
mm) PL. 11, FIG. 7.
Oliva spicata var. hemphilli Ford in Johnson, 1911.
Five syntypes ANSP 111697. One (H: 54.05 mm, D:
23.38 mm) is figured PL. 10, FIG. 8.
Oliva spicata var. perfecta Johnson, 1911. Three
syntypes ANSP 111729. One (H: 59.06 mm, D: 25.73
mm) is figured PL. 11, FIG. 5.
APEX 13(1-2): 1-61, 20 avr. 1998
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
Oliva rejecta Burch & Burch, 1962. One paratype
USNM 667317 (H: 37.24 mm, D: 16.01 mm) (see PL.
10, FIG. 10). One paratype at MNHN (H: 313, D:
13.6 mm) (see PL. 10, FIG. 9).
Oliva (Strephona) radix Petuch & Sargent, 1986.
Holotype USNM 841446 (H: 36.62mm, D: 22.51
mm) (see PL. 10, FIG. 6).
Oliva subangulata corteziana Petuch & Sargent, 1986.
Holotype USNM 841457 (H: 32.33 mm, D: 16.21
mm) (see PL. 10, FIG. 7).
Oliva spicata deynzerae Petuch & Sargent, 1986.
Oliva (Strephona) spicata deynzerae Petuch & Sargent,
1986. Holotype USNM 841452 (H: 53.19 mm, D:
21.95 mm) (see PL. 4, FIG. 5).
3. METHODS
3.1. Measurements.
The protoconch measurements NW, SPRO, MPRO,
LPRO, RESS, PATI7, PATI8 and the teleoconch
measurements PNW, H, L, LW, D, R, X, F, FG and
SUT that are utilized in this work have been defined in
detail by TURSCH & GERMAIN (1985, 1986, 1987).
They have been repeatedly tested and demonstrated to
be operational in the genus Oliva (see TURSCH,
GERMAIN & GREIFENEDER 1986a, 1986b; TURSCH &
HUART 1988; 1990; TURSCH 1988, 1994: TURSCH &
GREIFENEDER 1989a, 1989b; TURSCH, MissA &
BOUILLON 1992).
For a quick reminder, these measurements are
sketched in Fig. 2. Two important measurements do
not appear on this figure. NW is the number of nuclear
whorls and PNW is the number of postnuclear whorls.
Both are measured to 0.05 whorl. The way in which
Oliva shell measurements are best used is discussed in
TURSCH & GREIFENEDER (1996).
3.2. Analysis methods.
These are essentially the same as in TURSCH, MissA
& BOUILLON (1992) and TURSCH & GREIFENEDER
(1996). Each specimen can be represented by a point in
the attribute hyperspace, i.e., a space having as many
dimensions as there are variables (attributes) under
consideration. Any phenon will thus appear as a cloud
of points in the attribute hyperspace, the dimensions of
the cloud reflecting variability.
Separations (voids between clouds of points) in
hyperspace cannot be visualized directly. These voids
can be detected by special techniques of space
reduction, such as Principal Component Analysis
(PCA) or Factorial Discriminant Analysis (FDA). In
favourable situations, however, only two of the many
variables can be sufficient for evidencing the
separation of two or several groups. This can then be
represented on bidimensional graphs (scatter
diagrams). Such separations constitute conclusive
evidence: groups that are separated in two dimensions
6
are necessarily separated in the multidimensional
attnibute hyperspace. From our experience, patient
search for operational characters (the number of
possible combination of measurements can be very
large) will generally yield a clean-cut bidimensional
representation of the separations. The search for such
operational characters is greatly helped by preliminary
PCA and FDA studies.
Only scatter diagrams will be reported here: they
convey all the necessary evidence and are much more
heuristic than PCA or FDA graphs. Numerical results
will be reported here only when really needed.
Reporting, for instance, the very evident differences
separating ©. porphyria from ©. kaleontina in
numerical terms would be quite superfluous. It would
consume space and unnecessarily tax the patience of
the reader.
Reading graphs containing hundreds of points is
very strenuous. We have therefore preferred to
represent the distribution of attributes by plain
minimum convex polygons.
3.3. Interpretation of data.
As in nearly all taxonomic studies of Mollusks, the
morphospecies approach is adopted here. On the one
hand it is evident (but often overlooked) that one can
never give a real demonstration that two objects (in this
case, lots of shells) belong to the same morphospecies.
In practice, two specimens belong to the same
morphospecies 1f they are linked by an unbroken chain
of intermediate specimens (sympatric or not).
On the other hand, one can present convincing
arguments that two (or more) samples belong to
separate morphospecies. Two morphospecies are
distinct if their representative clouds in the attribute
hyperspace are separated by a void region: a
morphological gap. To prevent taxonomy from
becoming a game of chance, only full separations
(with no overlap) will be taken into account in this
work. If one cannot bring forwards such arguments,
then one has no choice: unseparated samples must be
considered as belonging to the same morphospecies
(awaiting further, objective arguments to the contrary).
Great care should be exercised in the interpretation
of the observed separations (see TURSCH &
GREIFENEDER 1996). The general problem of detecting
separations is much aggravated in multidimensional
hyperspaces (where one can get horribly lost). In the
bidimensional graphs used in this work, interpretation
is much safer but caution is neverteless required. First,
one has to consider the size of the samples. In scatter
diagrams, weak separations and separations obtained
on small samples (from our experience, V<6) might be
indicative but must be considered with caution. Small
gaps observed between small samples should be
interpreted with great prudence. All separations
accepted in this work are supported by additional
evidence: the observed gaps correspond to
discontinuities in the distribution of other characters,
unrelated to the nature of the axes of the graph (other
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
metric characters, colour pattern features, distributional
data, etc.).
In contrast, observed overlaps are always
significant, as these overlaps will persist if the size of
the sample is increased. Even one single specimen can
be legitimately tested by comparison with a large
sample. This allows the demonstration of overlaps in
the case of type material.
The interpretation of morphometric separations in
terms of distinct species is straightforward only for
samples that are syntopic (.e., actually live together, in
the same microbiotope). Only in these conditions can
morphological gaps be safely interpreted in terms of
reproductive barriers.
It has been shown (TURSCH 1994) that Oliva
species consist of a mosaic of distinct populations, each
being quite homogeneous. When a large enough
number of such local, conspecific populations are
compared, they invariably show considerable character
overlap. The species is thus represented by a
morphological continuum in the attribute hyperspace.
This is a set in which no population (or groups of
populations) can be separated from a// the others. Even
if two (or more) of the populations forming the
continuum can be easily separated from each other, the
gap is invariably bridged by another conspecific
population (or an unbroken chain of intermediate
populations). The boundaries of the morphological
continuum are the limits of the phenetic variability of a
species.
Completely separable populations belonging to the
same Oliva species can occur even within very short
geographic distances (TURSCH 1994). Separations
between pairs of conspecific populations are thus fully
expected (see MAYR & ASHLOCK 1991 and FUTUYMA
1986) and do not constitute grounds for specific
discrimination. Well on the contrary, observation that
very similar Oliva phena do never co-occur constitutes
a strong indication of their conspecificity (TURSCH
1995).
In the morphospecies approach, specific discrimination
can be established only by the separation of sets
containing as many different local populations as
possible. Even with a small number of specimens,
samples including Oliva shells from different localities
include much more of the total variability of the
species. Clear separations of such samples are more
probably significant.
Other problems can occur within the population level.
Sexual dimorphism can fortunately be neglected in the
genus Oliva. In contrast, some Oliva species have a
non-isometric growth (TURSCH 1997) and in such
instances, unless adequate precautions are taken,
artificial segregation of young and aged individuals can
easily occur.
r— D—+
Teleoconch, lateral view
MPRO
<< +
\
Protoconch,
apical view.
SPR
LPRO
L
SUT
(©)
Teleoconch,
apical view
*<— 7 —+
Protoconch,
lateral view
Fig. 2. Sketch of shell measurements used in this work.
APEX 13(1-2): 1-61, 20 avr. 1998
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
In theory, the objective approach advocated here
could either increase the number of species (by
detecting previously overlooked gaps) or reduce it (by
demonstrating previously overlooked overlaps). In
practice, it does lead to a sharp diminution of the
number of admitted species. This is frequently called
‘lumping' by collectors who ‘know’ their shells. Please
note that ‘splitters' and ‘lumpers' can exist only when
taxonomic decisions are a matter of personal opinion.
This can certainly be the case at the supraspecific level,
where one can disagree on where to ‘cut the branches’
of a phylogenetic tree. But the species is (or at least
should be) the most objective of all taxonomic
categories. At the species level, we should not even
have the choice between ‘splitting' and 'lumping”. These
two attitudes can, at best, be provisional strategies for
handling unsolved cases.
In summary, the method used in this work is very
simple. A large number of purposely unidentified
specimens are first shown to be separated into distinct
morphological groups. By careful selection of
characters (using PCA and FDA), the separations of the
groups could be demonstrated in bidimensional
representations. In spite of all efforts, these groups
could not be split any further (even when submitted to
PCA and FDA tests on all variables). We consider
these groups as separate morphospecies, a conclusion
supported by the fact that all additional specimens fall
into these clusters. If the type material of other taxa
cannot be separated from these groups, synonymy is
demonstrated
The species so defined can now be visually
identified with a rather high degree of confidence
because the distribution of morphometric characters is
correlated to the distribution of some (mostly
undescribed) visual clues. But these clues could be
fulgurator ,
complex À
25 /
polpasta
fulgurator
f. circinata
1:57
spicata complex —
Spicata
deynzerae
found only after morphometric analysis. Many
‘intuitive species’ erected only on other, uncorrelated
visual clues are listed in a specially long synonymy
(see Index to names).
4. RESULTS AND OBSERVATIONS
4.1. Species delimitation.
The inclusion of many additional specimens in the
morphometric analysis did not fundamentally modify
the conclusions previously drawn by TURSCH & HUART
(1990). For instance, all the Atlantic and Eastern
Pacific Oliva species are included in the scatter
diagram of Fig. 3, where many species are objectively
separated in one single operation. ©. foxi Stingley,
1984 (not considered in TURSCH & HUART 1990) is
completely separated from ©. spicata and ©. polpasta.
The species ©. incrassata, ©. julieta and ©. peruviana
(also not considered in TURSCH & HUART 1990) are
clearly separated from cognate species and will not be
treated here anymore, except for comparison.
AIT the Atlantic Oliva taxa still fall into four
objective groups, clearly separated even on one single
scatter diagram (see Fig. 4). One of these groups is ©.
flammulata Lamarck, 1811, confined to the West
African coast, with a subspecies ©. flammulata dolicha
Locard, 1896 in Cabo Verde (see TURSCH & HUART,
1988). It will not be treated here anymore, except for
comparison. The three others live in the warm waters
of the Western Atlantic. Two of these objective groups
correspond to the species ©. scripta Lamarck, 1811
and ©. sayana Ravenel, 1834. These groups are quite
homogeneous and could not be split into smaller,
objective units.
kaleontina
splendidula
foxi
peruviana
porphyria
flammulata
julieta
sayana
incrassata
undatella
Fig. 3. All Atlantic and Pacific species. Scatter diagram: RESS5 vs. NW. Minimum convex polygons. Every group is
well separated with other Variables, with the exception of the “O. fulgurator complex" not separated from the “O.
spicata complex”.
8
TURSCH, GREIFENEDER & HUART Oliva fulgurator and related species APEX 13(1-2): 1-61, 20 avr. 1998
SUT /(H-L)
0.25 +
scripta
\
reticularis- >
fulgurator SEULE N
flammulata
0.05 + À
LA 1 / (LPRO - SPRO)
L + + + t
1 2 3 4 5
Fig. 4. Separation of Atlantic Ofiva species. Scatter
diagram: SUT/H-L) vs. 1/(LPRO-SPRO). Minimum
convex polygons.
The third Western Atlantic group, the "O.
Julgurator-reticularis complex", has a much higher
variability, as attested by the relative dimensions of its
representative cloud of points in most dimensions of
the attribute hyperspace. Some of the populations that
it contains are indeed very dissimilar in aspect. In spite
of much effort spent to that end, this "complex" could
not be objectively split into smaller units. Inclusion of
additional specimens now established that ©. oblonga
Marrat, 1870, formerly thought by us to be a possible
candidate for separation (see TURSCH & HUART 1990),
does considerably overlap with the complex.
In the Eastern Pacific, all the separations previously
observed persisted, with only one exception. A scatter
diagram of R/L vs. NW/(LPRO-SPRO) (Fig. 5) clearly
separated ©. porphyria and ©. splendidula from two
large, distinct, but not homogeneous groups
These two unresolved groups (NH) were then
analysed in a scatter diagram of PATI8/NW vs. LPRO
(Fig. 6) yielding four groups corresponding to ©.
kaleontina, ©. undatella, ©. polpasta and the "O.
spicata complex". None of these final groups could be
objectively split into smaller units. One result differed
from the previous ones: with the inclusion of
additional, allopatric specimens, ©. polpasta
Duclos,1833 could not anymore be separated from ©.
kerstitchi da Motta, 1985 Jthis separation was
suggested in Tursch & Huart (1990) as a working
hypothesis awaiting further confirmation].
In spite of much effort to that end, no quantitative
discrimination of the "©. fulgurator-reticularis
complex" from the "O. spicata complex" could be
established, as already observed in TURSCH & HUART
(1990). À very large number of separation attempts
(including frequency histograms, scatter diagrams,
principal factor analysis, discriminant factorial analysis
and UPGMA clustering) invariably resulted in
considerable overlap between the two groups, as in the
example of Fig. 7. There is nothing really new in this
observation:
"The discrimination of... [O. melchersi and ©.
venulata] … from each other and from ©.
reticularis is a matter of extreme difficulty;
which Mr. Reeve escapes by uniting them all
together."
CARPENTER (1855: 464).
0.4 + RL PAT18 / NW
\ 0.6 +
NH D N spicata
Ts
NH _d
0.3 T 0.4+ kaleontina
\ polpasta
splendidula
undatella
porphyria 0.2 +
NW / (LPRO - SPRO) LPRO
0.2 . | | | | }
5 15 25 0.25 0.75 1.25
Fig. 5. Separation of Eastern Pacific Ojiva species.
Scatter diagram: R/L vs. NW/LPRO-SPRO). Minimum
convex polygons. The groups marked NH are not
homogeneous.
Fig. 6. Separation of Eastern Pacific Ofiva species.
Separations of the groups NH unresolved in Fig. 5.
Scatter diagram PAT18/NW vs. LPRO. Minimum
convex polygons.
APEX 13(1-2}) 1-61, 20 avr. 1998
0.7 :- X/R
Ps 4h SS
” Wy,_\
L ?e \
\ / > S
06! | Ke
/\ va
\ \ Se
| \ 7 Ts
| | \ A A
0 \ A PA
Cr 14
| ne E. PACIFIC 7
06 N LA
Re D/L
1LE | Il | | Ï
1.35 1:55 1:79 1.95
Fig. 7. Attempted separation of the "©. fulgurator-
reticularis complex" (marked "W. Atlantic") from the "O.
spicata complex" (marked "E. Pacific"). One example
amongst many. Scatter diagram: X/R vs. D/L. Minimum
convex polygons.
Because of their long-standing disjunct
geographical distribution (and only for that reason),
this constant overlap does not necessarily mean that the
two groups are of the same species (see $ 5.4).
Many of the groups defined here above have quite
characteristic protoconchs. Examples are given in Fig.
8 and Fig. 9. One will note the exceptionally great
variation range of the protoconchs of ©. fulgurator and
O. spicata. This variation is way larger than for any of
the Indo-Pacific species that we have examined. The
ranges of variation of all subsamples form an unbroken
continuum (see $ 4.4).
Two characteristics of the shell colour pattern
confirm the close (and expected) relationship of the
Western Atlantic and Eastern Pacific Oliva faunas, and
set a number of species apart from all others. In the
genus Oliva, the presence on the shell of a peculiar
colour pattern located just under the filament channel is
a reliable indication that one is dealing with an
American species (for details on the filament channel,
see VAN OSSELAER & TURSCH 1994). This repeated
pattern consists in bundles of divaricate, fasciculated,
fine lines meeting at a central point or a small blotch,
right at the sharp exterior edge of the channel (see Fig.
10). It is -as expected- very variable but nevertheless
highly recognisable. It will be referred here under as
the "fasciculated pattern", for short. The presence on
the shell of this pattern, quite unique in the genus
Oliva, is a safe indication that one is dealing with either
a Western or an Eastern American species. Ît is indeed
present in all American Oliva species, except O.
kaleontina, ©. peruviana and ©. undatella (see Plate
1). It is not clearly present in ©. flammulata Lam.
1811 (possible intermediates are occasionally met). It
is not found in any Indo-Pacific species (with the
10
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
possible exception of an intermediate pattern found on
the Hawaïian, deep water ©. richerti Kay, 1979 from
Hawaï). The presence of this same subchannel pattern
in species as different as ©. porphyria and ©. spicata
could be interpreted as indicating a common, but
ancient ancestry.
There is one other shared feature in the colour
pattern. Most species of the genus Oliva display two
more or less diffuse spiral bands of darker colour on
the body whorl. It has been shown by GREIFENEDER
(1984) that the relative position of these bands in many
American Oliva species is consistently lower than that
observed in their Indo-Pacific congenerics.
Another argument yet could be found in the
orientation of the shell microcrystalline layers. The
distribution of angular values in both the Western
Atlantic and Panamic Oliva species has been shown to
differ from that of their Indo-Pacific counterparts. This
should be interpreted with caution because the
phenomenon could possibly be of adaptative nature
(TURSCH & MACHBAETE 1995).
4.2. Homogeneous, distinct local populations.
Except for occasional colour variants, the local
populations of the members of the “/w/gurator-
reticularis complex” are quite homogeneous in shape
and general outline (see DIAZ & PUYANA 1994: 202).
Albeit variable (colour polymorphism results in
occasional colour variants such as darker or paler
individuals) the ground colour of the shell is generally
cryptic, matching the substrate, just as for most Indo-
Pacific Oliva species (see VAN OSSELAER ef al. 1993).
This homogeneity (probably due to the inefficiency of
immigration into generally large populations —-see $
6.1- and possibly maintained by selection) does restrict
even more the intra-population variation.
In contrast, inter-population variation is much
greater, considerable differences being often observed
between specimens from different localities. This was
already clearly perceived by DIAZ & PUYANA (1994:
202), who wrote about ©. bewleyi Marrat, 1870:
“Note: this species is very variable in its colour
pattern, even within one local population.
Variations in the form of the shell (height of the
spire, ratio height/width, etc.) are frequent
between populations but not so within one same
population. Within the abundant material of ©.
bewleyi from different localities of the
Colombian and Venezuelan Caribbean
examined by us, and applying the criteria of
Sargent & Petuch (1986), not less than six
different 'species' or ‘subspecies' exist in the
region.”
(translation ours).
That the distribution of most of the populations is
quite restricted, is evidenced by the fact that the
experienced collector can often guess the origin of a
given specimen.
TURSCH, GREIFENEDER & HUART Oliva fulgurator and related species APEX 13(1-2): 1-61, 20 avr. 1998
Les O. fulgurator ©. fulgurator
circinata
20 2.70
2.30 225 j
1 2 3 4 9
bi 1 k— 1 1 E—— k— —
0.36 - 0.90 - 1.46 0.45 - 1.06 - 1.73 0.41-0.77-1.11 0.19- 0.62 - 0.99 0.36 - 0.74 - 1.03
3.30
CE 2.80
2.70 275
À | —.
FH — b— 4 1 k—— PRE"
0.27 - 0.62 - 0.96 0.27 - 0.63 - 0.98 0.33 - 0.64 - 1.03 0.26 - 0.59 - 0.84 0.36 - 0.70 - 0.98
©. spicata O. spicata
deynzerae
2.85
3.35 2.35 2.25 2.35
11 12 13 14 a
es | + + 1 h—-——
0.30- 0.66- 1.03 0.24- 0.50- 0.75 0.22- 0.63- 1.06 0.29 - 0.73 - 1.17 0.42 - 0.77 - 1.02
3.00
3.05 D 2.55
245
: : nr N ,
+ + + +---—1 b———
0.21 -0.51-0.87 0.31 - 0.66 - 0.99 0.29 - 0.62 - 0.98 0.27- 0.53- 0.88 0.39- 0.75- 1.00
O. polpasta ©. scripta
3.10
3.00 3.20
23 24
7
0.37 - 0.62 - 0.89 0.23 - 0.49 - 0.72 0.29 - 0.56 - 0.82 0.30-0.62-0.81
Fig. 8. Protoconchs (Atlantic). Scale bars: 1 mm. Figures to the left of apex: number of nuclear whorls (NW).
Figures below scale bar: measurements SPRO, MPRO and LPRO of first protoconch volutions (see Fig. 2).
1-10: O. fulgurator (Rôding, 1798). 1: Isla Boracha, VENEZUELA (BT-0984); 2: ARUBA (BT-4964), 3: BaHamas (BT-
3418), 4 Guantanamo, CUBA (BT-2668),; 5: Lac Bay, BONAIRE (BT-3872); 6: Guantanamo, CUBA (BT-2669); 7: Lac
Bay, BONAIRE (BT-3874); 8: Brevard County, FLoRIDA (BT-2764). 9-10: ©. fulgurator forma circinata Marrat, 1871.
9: Alagoas, BRaziL (BT-2114), 10: Rio de Janeiro, BRaziL (BT-2113). 11-18: O. spicata (Rôding, 1798).
11: Baja California, MExIco (AB-b207), 12: Cebaco |., PANAMA (BT-3755); 13: Baja California, Mexico (BT-4289);
14: Guerrero, Mexico (BT-5335), 15: Baja California, Mexico (BT-0346), 76: Baja California, MExXIco (RF-b06);
17: Baja California, MExIco (BT-4123); 18 Baja California, MExIco (AB-b384). 19-20: O.spicata deynzerae Petuch
& Sargent, 1986. 19: Cocos Is., CosTA Rica (BT-5447) 20: Cocos Is., CosTA Rica (BT-5446).
21-22: O. polpasta Duclos, 1840. 21: Oaxaca, Mexico (BT-4613, forma kerstitchi), 22: Cebaco |., PANAMA (BT-
3781). 23-24: O. scripta Lamarck, 1811. 23: HonbuRraAS (BT-2756); 24: Hairi (BT-2379).
11
APEX 13(1-2): 1-61, 20 avr. 1998 Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
©. peruviana | ©. incrassata
4 00 SN, 4.20 =
A — | om dé N Ë 3.60 \
LT ES PER NE |2 340 3 té
E— 1 \ | b-— \ A À gene
0.14-0.26-0.42 } 0.17-0.29-0.43 \ 0.25 - 0.38 - 0.58 0.29 - 0.48 - 0.68
F+— - - +
| O. foxi O. julieta
| 4.80 CN SOA
ETUIS pe PT ere 8
/ |
SE, —
1 Rd À
F1 \Lb 5 Et Par +
| 0.16 - 0.29 - 0.44 \ 0.18 - 0.31 - 0.49 / 0.17 - 0.29 - 0.48 0.20- 0.32 - 0.45
| O. kaleontina FVÉRS O. porphyria
ne _ fes \ 4.75
PÉ DL, =>
\9 RP di 11 12
LS EE M Le /
| Sr \
+ Re h
) - A \ RE: —
0.09 - 0.18 - 0.31 0.08 - 0.18 - 0.30 0.20 - 0.38 - 0.58 0.19 - 0.40 - 0.57
©. splendidula ©. undatella
BTS SN DLOD 525 Le
AN fe \ 14 15 | 16
ne ph
/ \
| + =
F4 +1 4 + NZ + 1 \
l 0.09 - 0.19 - 0.35 0.09-0.18-0.32 0.13 - 0.21 - 0.30 0.08 - 0.16 - 0.29
©. flammulata ©. sayana
450 17 440 _— 18 3:90 19 3.90 20
= |, oi = +
0.10 - 0.20 - 0.35 / 0.10-0.21-0.35 0.19 - 0.32 - 0.48 0.16 - 0.32 - 0.49
Fig. 9. Protoconchs (Eastern Pacific). Scale bars: 1 mm. Figures to the left of apex: number of nuclear whorls
(NW). Figures below scale bar: measurements SPRO, MPRO and LPRO of first protoconch volutions (see Fig. 2).
1-2: ©. peruviana Lamarck, 1811. 1: Iquique, CHILE (BT-5785), 2: iquique, CHILE (BT-5784). 3-4: O. incrassata
(Lightfoot in Solander, 1786). 3: Puerto Penasco, MExIco (DG-4224/3),4: Michoacan, MExIico (DG-4224/2).
5-6: O. foxi Stingley, 1984. 5: Cocos Is., CosTA Rica (BT-3326), 6: Cocos Is., CosTA Rica (BP-b07).
7-8: O. julieta Duclos, 1840. 7: Michoacan, Mexico (DG-4224/3); 8: Michoacan, Mexico (DG-4224/3).
9-10: O. kaleontina Duclos, 1835. 9: Cebaco |., PANAMA (BT-3751), 10: St. James |., GALAPAGOS (BT-4275),.
11-12: O. porphyria (Rôding, 1798). 11: Gubernadora |., PANAMA (BT-4464); 12: Sonora, MExIco (BT-0346).
12-13: O. splendidula Sowerby, 1825. 13: Perlas Is., PANAMA (BT-3731), 14: Manzanillo, MExico (BT-4023).
15-16: O. undatella Lamarck, 1811. 75: Colima, Mexico (BT-0331), 16: Venado I., PANAMA (BT-1666).
17-18: O. flammulata Lamarck, 1811. 17: Port Gentil, GABON (BT-2087), 18: Gorée, SÉNÉGAL (BT-2127).
19-20: O. sayana Ravenel, 1834. 19: Indian River, FLORIDA (BT-4064), 20: Marco Beach, FLORIDA (BT-3108).
12
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
Le
edge of filament channel
Fig. 10. Some examples of "fasciculated" subchannel colour markings (see & 4.1).
Although some general geographical trends can be
noted (see $ 4.5), neighbouring populations do not
necessarily resemble each other very much The
general distribution of many single characters shows
chaotic interdeme variation [like the “crazy quilf
distribution observed for Cerion by GOULD &
WoOODRUFF (1978)] and would seem to fairly reflect
habitat discontinuities.
4.3. Systematic allotopy.
4.3.1. The “O. fulgurator-reticularis complex”
Much time has been spent in interviewing many
experienced local collectors (ranging from professional
malacologists to fishermen collecting molluscs for the
shell trade). All were unanimous in reporting that
within the “fulgurator-reticularis complex”, shells of
different forms ('species' ?) are never found living
together. This has also been our personal experience
during collecting trips in Brazil, Colombia, Mexico,
Venezuela and the Virgin Islands. The populations of
shells with different forms are invariably a/lotopic.
It is important to stress the restricted meaning of
“living together”. One could indeed object that some
distinct populations can live within short distance (for
instance, at least two forms have been reported from
Gonave, Haïti, by PETUCH 1986, and at least three
different forms live in Aruba. Detailed field studies in
the Southern Pacific have demonstrated that
completely distinct, conspecific populations of several
species have been observed to be separated by only a
few hundred meters (VAN OSSELAER ef al. 1993). The
scale of sympatry in the genus Oliva is thus much
smaller than generally thought (TURSCH 1994) and it
should, for safety, be reduced to the scale of syntopy
(i.e. found living in the same microbiotope, within a
distance of meters).
4.3.2. The “O. spicata complex”
From all the information we could gather, the same
situation is met for the “O. spicata complex” in the
Eastern Pacific. The only exception known to us 1s the
report by BURCH & BURCH (1962) on the coexistence
of the "species" ©. rejecta Burch & Burch, 1962 (see
Section 7, Systematics, under ©. spicata) with ©.
venulata Lamarck, 1811. The authors wrote: "Both
forms are found on the same tide flats at La Paz, Baja
California, Mexico, with no intergrades in many
hundreds of specimens". One will notice the use of the
word "forms" to designate what the authors consider
distinct species. Nevertheless, this had to be taken
seriously because, if this coexistence were factual, it
could falsify the very premises upon which the
conclusions of the present work do rest.
One of us (BT) recently went to La Paz for an in
situ check of the situation. Large numbers of live
specimens were observed at eleven tide flats in Bahia
La Paz (see map, PI. 2) (e.g. 78 specimens at Punta de
Leon, 129 at Herendira, 318 at Balandra). The previous
observations made in the Caribbean and in the South
Pacific were fully verified: all the Oliva spicata
populations that were seen were local morphs, with
cryptic colouration (matching that of the substrate), and
quite homogeneous in characters. No form absolutely
identical to the "real" ©. rejecta was not found
(although the Herendira population is quite similar, see
PL 2). It might live in another of the very numerous
isolated bays of the region (their systematical study
would take very considerable time). Or its habitat may
have been destroyed (the original beaches of La Paz
City are said to have been washed away after the
construction of the Malecôn, many years ago; the sand
of the present beaches is imported).
Around La Paz, Oliva spicata ïis collected
(overcollected?) in large quantities for the tourist trade.
So the conclusions from our field observations were
fully confirmed by the examination of well over 100
kilos of specimens in the possession of local fishermen
(one single person had a lot of 60 kilos -estimated at
over 8000 specimens). None of the local collectors
(some gathering ©. spicata for over 25 years) could
remember a single case of co-occurrence of different
forms.
It would thus seem that BURCH & BURCH were
simply given incorrect locality data (as it only too often
happens for commercial specimens) and that the case
of O. rejecta cannot be construed as a counter-example.
13
APEX 13(1-2): 1-61, 20 avr. 1998
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
D/L
0.65 |
0.60 |- :
0.55 +
0.50 +
175
Fig. 11. Example of total separation of selected local
samples within the “O. fulgurator-reticularis complex”
(gray zone). Scatter diagram: XR vs. D/L . Minimum
convex polygons. 1. CURAÇAO, (O. cfr. nivosa Marrat);
2. PUERTO Rico (©. reticularis Lamarck), 3. BARBADOS,
deep water (O0. barbadensis Petuch & Sargent);
4. BERMUDA (0. bifasciata jenseni Petuch & Sargent);
5. BRAZIL, Bahia (O. circinata Marrat), 6. VENEZUELA,
Margarita |. [O. fulgurator (Rôding)].
4.4. Morphological continuum.
Much effort (research on this project started in 1987)
has been invested in trying to break the “O. fulgurator-
reticularis complex” and the “O. spicata complex” into
objective subgroups, mainly by using the methods
already applied to the Indo-Pacific "©. miniacea
complex" based upon ©. miniacea (Rôding, 1798)
(TURSCH & GREIFENEDER 1996). All these attempts
(many combinations of variables were tested) failed
consistently.
In the Western Atlantic, all the local populations of
the “©. fulgurator-reticularis complex” can be
objectively separated from some (or even many)
others, as illustrated by the scatter diagrams of Figs. 11
and 12. The same situation is met in the Eastern Pacific
for the members of the “O. spicata complex”, as
illustrated in the scatter diagrams of Fig. 13.
These separations are invariably obliterated by the
inclusion of some additional population(s). As one
example amongst many, Fig. 14 groups the populations
previously separated in Figs. 11 and 12, with the
addition of only four other local samples. Every one of
these distinct populations is now linked to every other
by an unbroken chain of intermediates, forming a
continuum.
Even without any measurement, the existence of a
continuum can be sensed just by glancing at Plates 6, 7,
8 and 9, in which the type material of the various taxa
constituting the “O. fulgurator-reticularis complex”
14
has been arranged by grouping specimens according to
resemblance. These plates, however, give only a
limited view of the total variation range because many
local forms have not been described.
0.60
0.55
0.50
1.35 1.55 1.75 1.95
Fig. 12. Example of total separation of selected local
samples within the “O. fulgurator-reticularis complex”
(gray zone). Scatter diagram: XR vs. D/L (same
variables as in Fig. 11). Minimum convex polygons.
7. CUBA, Guantanamo (unnamed form); 8. ST. MARTIN
(O. nivosa Marrat), 9. FLORIDA, off Punta Vadra, deep
water (O. bollingi Clench), 10. CoLOMBIA, Santa Marta
(O. goajira Petuch & Sargent): 11. VENEZUELA,
Paranagué Peninsula [O. cfr. fulgurator (Rôding)]].
0.5 0.6 0.7 0.8 0.9 1.0
Fig. 13. Example of total separation of selected local
samples within the “O. spicata complex” (gray zone).
Scatter diagram: MPRO vs. R/Pnw. Minimum convex
polygons. 1. PANAMA, Bayarena, 20-30 m. 2. MEXICO,
Baja California, La Paz, shallow water; 3. Cocos Is., 12-
20 m (O. spicata deynzerae Petuch & Sargent).
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
0.65
0.60
0.55
0.50
1.35
1.55 1.75 1:95
Fig. 14. Local populations within the “fulgurator-
reticularis complex” (gray zone) form a morphological
continuum: an example. Same variables and
populations as in Figs. 11 and 12, 4 populations added:
12 to 15. Scatter diagram: XR vs. D/L). Minimum
convex polygons. 1. CURAGÇAO, (O. cfr. nivosa Marrat),
2. PUERTO Rico (0. reticularis Lamarck); 3. BARBADOS,
deep water (O. barbadensis Petuch & Sargent);
4. BERMUDA (©. bifasciata jenseni Petuch & Sargent);
5. BRAZIL, Bahia (O. circinata Marrat);, 6. VENEZUELA,
Margarita |. [O. fulgurator (Rôding)]; 7. CUBA,
Guantanamo (unnamed form), 8. ST. MARTIN (O. nivosa
Marrat), 9. FLORIDA, off Punta Vadra, deep water (O.
bollingi Clench); 10. CoLomBiA, Santa Marta (O. goajira
Petuch & Sargent): 11. VENEZUELA, Paranagué
Peninsula [O. fulgurator (Rôding) variant];
12. DOMINICAN REPUBLIC (©. jamaicensis zombia Petuch
& Sargent), 13. HONDURAS (0. aff. ernesti Petuch);
14. ARUBA (O. cfr. sargenti Petuch), 15. ST. VINCENT
(unamed, dark form).
Petuch & Sargent (1986: 119-122) claimed that,
"based on shell morphology", the taxa we here place in
the “O. fulgurator-reticularis complex” break up into
groups: the reticularis group ("characterized as being
ovate shells with rounded outlines and by having
variable amounts of triangle net color patterns"), the
bifasciata group ("slender, cylindrical, elongated shells
with fairly straight sides"), the nivosa group
(‘elongated shells with straight or slightly rounded
sides … flattened spires and intricate, fine-netted color
patterns") and the fisiphona group (said to “
resemble the Panamic ©. spicata and ©. incrassata
groups"). We could find no basis whatsoever on which
to segregate the above groups.
Exactly the same situation was observed for the “O.
spicata complex”. As one example amongst many, Fig.
15 adds just two other local samples to the populations
previously separated in Fig. 13. Here again, one sees
the formation of a continuum.
In conclusion: we have found no way of separating
any of the populations (or group of populations) from
all the others. The “O. fulgurator-reticularis complex”
and the “©. spicata complex” each consist of a
morphological continuum of local forms (see $ 3.3).
4.5. Clinal variation.
Although neighbouring populations do not necessarily
resemble each other (see $ 4.2), the populations of the
“O. fulgurator-reticularis complex” display a quasi-
clinal variation of some characters, such as the
protoconch size. This can be seen, for instance, in Fig.
16, showing the distribution of the largest local
measurements of PATI8 (the diameter of the
penultimate nuclear volution, see Fig. 2).
This quasi-clinal variation, from Northern South
America outwards, is not restricted to protoconch
characters but is also reflected in the general shape of
shells. Globose shells (©. fulgurator s.s.) are found
only around Venezuela and Aruba, whereas more
cylindrical, elongated shells are found at both ends of
the distribution range, in Brazil (the form circinata)
and in Florida. Here again, the variation is roughly
clinal, as shown in Fig. 17, the distribution of the
largest local measurements of D/L (breadth of the body
whorl relative to the length of the lip). A quite similar
clinal distribution (not illustrated here) is observed for
the mean values of D/H.
No such clines have yet been observed for the
forms of the “O. spicata complex”, in the Eastern
Pacific.
R/PNW
3.0
2.5
2:01
Toi
LE MPRO
1.0 |
0.5 0.6 0.7 0.8 0.9 1.0
Fig. 15. Local variants within the “O. spicata complex”
(gray zone) form a morphological continuum: an
example. Same variables and populations as in Fig. 13,
2 populations added: 4 and 5. Scatter diagram: XR vs.
D/L). Minimum convex polygons. 1. PANAMA, Bayarena,
20-30 m. 2. MEXIco, Baja California, La Paz, shallow
water; 3. Cocos Is., 12-20 m (O. spicata deynzerae
Petuch & Sargent). 4. MExico, Baja California, Santa
Rosalia, shallow water; 5. PANAMA, Cebaco I. 35-40 m.
15
APEX 13(1-2): 1-61, 20 avr. 1998 Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
ES < 1.20
1.20 - 1.39
KW 1.40 - 1.59
BE :60-:7°
ES - 50
Fig. 16. "O. fulgurator-reticularis complex". Clinal variation of the protoconch measurement PAT18. Distribution of
maximal observed values (see text $ 4.5). Only a few values are given, for clarity. The values for isophene lines are
PS
arbitrary.
SD |
c)
[ D/L= 0.44 D/L=055 D/L=0.59 D/L= 0.62
) (0.44 - 0.55
Fig. 17. "O. fulgurator-reticularis complex". Clinal variation of the teleoconch character D/L. Distribution of maximal
observed values (see text $ 4.5). Only a few values are given, for clarity. The values for isophene lines are
arbitrary.
16
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
| PAT16
0.65
O. spicata complex
0.45 -—
©. spicata
deynzerae
GES , (LPRO-SPRO)
=) Il
0.45 0.65 0.85
Fig. 18. "O. spicata complex". Separation of O. spicata
deynzerae. Scatter diagram: PAT16 vs. (LPRO-SPRO).
Minimum convex polygons. See text $ 5.3.
5. INTERPRETATION
5.1. Oliva fulgurator (Rüding, 1798).
Three independent arguments indicate that all the
cognate taxa forming the “©. fulgurator-reticularis
complex” constitute one single, highly variable species.
We are certainly not the first to reach this conclusion:
“OL. fusiformis, Lam. [the former name for ©.
Julgurator (Rôding)], is a shell that appears
characteristical when one has only fypes, but
one finds all possible intermediates and we do
not understand why Mr. Reeve did not propose
its reunion with rericularis, as he has done for
others."
DUCROS DE SAINT GERMAIN (1857: 54)
(translation ours)
The first argument is purely phenetic. In spite of
much effort to that end, none of the taxa constituting
the “OC. fulgurator-reticularis complex” (cited in the
synonymy of ©. fulgurator) could be separated from
all the others on the basis of shell measurements. The
range of variation of both the teleoconchs and the
protoconchs is quite extraordinary for the genus Oliva,
but all intermediate forms are present. The various
populations form a morphological continuum (see $
4.4). Analysis of the colour patterns of the shells fully
confirms this view.
The second argument is based on distribution data.
All the taxa constituting the "©. fulgurator-reticularis
complex" are systematically allotopic (see $ 4.3). This
mutual exclusion of so many Oliva 'species' is highly
unusual. It is well known to anyone who has collected
in the Indo-Pacific that most (if not all) species of
Oliva are found together with other, congeneric
species. Up to 12 species of Oliva have been observed
to live together in the same microhabitat in Hansa Bay,
Papua New Guinea (VAN OSSELAER ef al. 1993).
PETUCH & SARGENT (1986) report that “over twenty”
Oliva species can be found living together in some
Indo-Pacific localities. So, one must now ask the
question: “How can we explain that the many
Caribbean ‘species’ of the ‘©. fulgurator-reticularis
complex’ do never occur together?” The simplest (and
most likely) answer is that they are not different
species but only local populations of the same species.
À third, independent argument for the
conspecificity of all the populations forming the “O.
Julgurator-reticularis complex” stems from the quasi-
clinal variation of some shell characters over the
distribution area (see $ 4.5). It is most improbable that
the observed pattern of variation reflects an
environmental cline. The 1.20-1.39 mm isophene zone
in Fig. 16, for instance, stretches over very varied types
of sediment, in very diverse physical conditions. An
interpretation in terms of clinal gene variation within a
same species is much more likely. Note: the observed
pattern (of central populations being distinct from
Northern and Southern populations which resemble
each other) appears to be rather common in other
zoological groups. It is familiar to ornithologists under
the name of "leapfrog" pattern of geographic variation
(see REMSEN 1984).
The differences in protoconch size observed
between extreme forms of the "complex" are much
greater than in any other Oliva species we know of.
This does not indicate separate specific status, the
range of variation being continuous.
Many of the populations forming the species are
today commonly referred to as ©. reticularis Lamarck,
1811, although the original concept of the species was
propably quite different (see the section Systematics).
We are nevertheless compelled to use the name ©.
fulgurator (Rôding, 1798), which has priority,
notwithstanding that it originally designated and is still
commonly used for a form which has a restricted
distribution and is less common in collections.
5.2. Oliva spicata (Rôüding, 1798).
As in the case of ©. fulgurator (Rôding, 1798), all the
examined populations of the "O. spicata complex"
(save one, see $ 5.3) form one single, compact
morphological continuum (see $ 4.4) We have
therefore to consider these populations as members of
one same species: Oliva spicata (Rôding, 1798). Here
also, the range of variation of both the shells and the
protoconchs is quite extraordinary for the genus Oliva,
but all forms are linked by intermediates. Analysis of
the colour patterns of the shells does again fully
confirms this view.
5.3. Oliva spicata deynzerae Petuch and
Sargent, 1986.
This taxon (see PL. 4, FIG. 5) is restricted to the Cocos
Islands and appears to be very closely related to Oliva
spicata (Rôding, 1798). It is nevertheless objectively
separated from al! other forms of the "O. spicata
complex" in a scatter diagram of LPRO-SPRO vs.
PATI6 (see Fig. 18). Very similar results are obtained
1
APEX 13(1-2)}: 1-61, 20 avr. 1998
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
with scatter diagrams of SUT/L vs. PAT16 and of NW
vs. PATI6 (not illustrated), the major discriminant
being PATI6.
The separation gap is narrow and rests upon a
rather small sample (7 only specimens of the Cocos Is.
taxon), so it is not impossible that the observed gap
could be filled by additional specimens. We shall
follow the advise of MAYR & ASQUITH (1991: 37) on
such cases (allospecies) and attach the Cocos Is.
population as a subspecies (deynzerae Petuch and
Sargent, 1986) to its closest relative, ©. spicata
(Rôding, 1798).
5.4. O. spicata and ©. fulgurator: separate
species ?
It has been seen ($ 4.1, fig. 7) that the Eastern Pacific
“O. spicata complex”, as a whole, could not be
objectively separated on morphometrical grounds from
the Western Atlantic “O. fulgurator-reticularis
complex”. Were it not for the existence of the Panama
land bridge, one would have little choice but to
combine the two complexes into one single
morphospecies. This logical step was indeed taken long
ago by DUCROS DE SAINT GERMAIN (1857: 52-56), at a
time when the locality data of most shells were
unreliable. His ©. reticularis Lamarck, 1811 -which he
considered to have à nearly world-wide distribution—
included a long list of taxa, known today to be
restricted either to the Eastern Pacific or the Western
Atlantic faunas.
Within the biological species concept, we cannot
anymore take such a simple stand (and we find
ourselves in the general problem of species with a
discontinuous distribution). The two "complexes"
cannot anymore meet to possibly interbreed, and have
now been separated by the Panama land barrnier for an
estimated 1.6 million years (see PETUCH & SARGENT
1986: 119). Therefore, as a working hypothesis until
tested by genetic studies, the “O. fulgurator-reticularis
complex” and the “©. spicata complex” will be here
considered as distinct species.
The two complexes certainly share a common
ancestry and constitute one more example of the many
pairs of ‘geminate species', sister taxa of which one
element is present in the Panamic region, the other in
the Caribbean. In nearly all cases, these ‘sister taxa’ are
considered to now form distinct species. Final closure
of the waterway linking the Pacific to the Atlantic
occured about 1.6 Ma (millions years ago) according to
PETUCH & SARGENT (1986: 119), 3.1-2.8 Ma according
to COATES & OBANDO (1996: 21), in any case an
acceptable "divergence time " for species separation
(see COLLINS 1996). Today, the Caribbean and the
Panamic regions are not only geographically separated;
they constitute two strikingly different realms.
Southern Caribbean waters are in average 2° C warmer
and 1.5 ‘% more saline than those of the eastern
equatorial Pacific. The latter, in addition, have strong
seasonal upwellings causing large increases in primary
productivity (see TERANES, GEARY & BEMIS 1996).
18
The eastern equatorial Pacific also has much stronger
tides, is subject to El Niño southern oscillation climate
anomalies, more intense predation and a much poorer
coral reef development (see JACKSON, JUNG &
FORTUNATO 1996).
Populations of the “©. /fulgurator-reticularis
complex” (Western Atlantic) often differ amongst
themselves as much as they differ from populations of
the “O. spicata complex” (Eastern Pacific). But the two
"complexes", although not objectively separable by
shell measurements, do nevertheless have different
trends. In general, Panamic specimens have a more
puncticulated colour pattern on the body whorl (the
starting point of the chevrons is marked by a dark
spot). In addition, the body whorl is often more angular
at the lip (this is not detectable by our measurements),
the "hifasciata colour pattern" with two sharply
contrasting dark, spiral bands (see PI. 99, Fig. 99) is
absent (or at least very rare), while all dark
(melanistic?) specimens are much more frequent.
So, albeit a few cases could lead to confusion, the
experienced Oliva student does today rarely err in
separating Atlantic from Panamic specimens. In most
instances, it is quite possible that we just recognise
well-known localities, characterized by familiar forms,
instead of using reliable, objective discriminants at the
species level. Errors in separation were much more
common one century ago. Many kinds of shells were
then available (possibly more than today, due to the
variety of ports of call of the sailing ships) but their
origin was uncertain (see $ 1.1.2) and no reliable
conclusions could be drawn.
Taxonomic distinction of non objectively separable
taxa of on the basis of evident geographic isolation is
not restricted to taxa separated by the Central American
land barrier and is now widely admitted. For instance,
the Panamic Conus dalli Stearns, 1873 and the Indo-
Pacific C. textile L., 1758 are not separable (according
to À. KOHN, private communication 1997).
6. DISCUSSION
6.1. Partial isolation of populations.
The most striking characteristic of the "©. fulgurator-
reticularis complex" and the "O. spicata complex" is
the high endemism of mutually exclusive local forms
(many of these forms have been named). This has also
been recognized by others. For instance, Petuch &
Sargent (1986: 120), refering to their "hifasciata
group" (see $ 4. 4) say that "Many of these species are
endemic to isolated seamounts and deep water
pockets".
The situation is far from being unique in the genus
Oliva. Within the distribution limits of the species, and
provided the species is present, every isolated little
beach of the Indo-Pacific that we have examined
appears to have one (and only one) local form of ©.
oliva (L., 1758). Many of these local forms are quite
recognisable. Of course, intrapopulation variants are
TURSCH, GREIFENEDER & HUART
common, but these are linked by sympatric (syntopic)
intergrades. This is the best argument for saying that ©.
oliva is only one species (all populations forming one
single morphological continuum). Very similar cases
are provided by ©. miniacea (Rôding, 1798), O.
amethystina (Rôding, 1798) and many other Oliva
species. We are just lucky that many of these local
Indo-Pacific forms have not yet been named as species
or subspecies.
One possible explanation is genetic. Such a
situation can occur if successful, long-distance
transport and settlement of the larvae is frequent
enough to allow occasional gene flow between
conspecific populations but rare enough to allow some
genetic drift of the local isolates. The efficiency of
larval transport and settlement will of course vary from
species to species, 1f it is highly efficient there will be
little local variation. Many Oliva species [for instance
O. splendidula, ©. porphyria, ©. caerulea (Rôding,
1798)] are indeed very constant over all their
distribution range.
Another explanation would be to consider that the
different local populations are only ecotypes, anising
from the expression of a same genotype in different
local environments. In this hypothesis, larval transport
should play a secondary role, if any. The great number
of local forms within many Oliva species would also
imply an extraordinary sensitivity to minute
environmental changes.
Distribution data support the first, genetic
hypothesis. No Oliva species has been reported from
Clipperton Island (EMERSON, 1994), or from the
Revillagigedo Islands (EMERSON, 1995), distant of
about 600 and 375 miles from Mexican mainland,
respectively. The much larger Galapagos (see FINET
1991) -about 600 miles from the mainland-— have two
species [O. porphyria (Linnaeus, 1758) and ©.
kaleontina Duclos, none belonging to the “O. spicata
complex”]. The somewhat less isolated Cocos islands
(about 300 miles from the mainland) have one endemic
species (©. foxi Stingley, 1984) and one endemic
subspecies (©. spicata deynzerae Petuch & Sargent,
time
Sp.A
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
1986). This indicates a weak efficiency of successful,
long-distance transport and settlement of larvae in the
“O. spicata complex”. Established populations
generally contain large numbers of individuals, so the
effect of ocasional larvae immigration can be expected
to be rapidly diluted in a large gene pool.
Dr. P. E. PENCHASZADEH (private communication)
together with Dr. Juan DIAZ, observed and collected in
small islands off Venezuela several specimens of a
form of “Oliva circinata” laying long ribbons of eggs,
anchored in the substrate. This phenomenon (not
reported yet for other Oliva species) could explain the
great endemism of local forms, larval development of
Oliva being known to occur within the egg capsule
(OLSSON & CROVO 1968).
The "crazy-quilt" distribution pattern of local
populations is not unique to Oliva species. A similar
pattern is observed for many taxa of Conus in the Cabo
Verde region and for many forms of Cymbiola
(Cymbiolacca) pulchra in Quensland, Australia (POPPE
& GoTo, 1992). Stable differences between allopatric
populations were reported by PARTH (1995) for taxa of
the muricid genus Haustellum .
6.2. Dynamic environment.
One can quite safely assume that speciation in the
genus Oliva follows the normal, allopatric (or
vicariant) process (see Fig. 19, a, b). This starts by the
advent of a geographical barrier restricting the gene
flow between populations of a same species. These
populations (now partially or totally isolated) will then
undergo divergent evolution, possibly to the point
where they cannot interbreed any more when brought
back into contact. Once started, the speciation process
is not irreversible. It might abort for a variety of causes
(see Fig. 19, c), for instance if the geographical barrier
does not persist long enough (which is precisely the
case advocated here above). In any case, speciation 1s
not a sudden event. The process takes place over a
large number of generations and there is necessarily a
period in time during which the speciation issue is
undecided.
. Speciation
issue settied
. Speciation
issue undecided
advent of
external barrier
variation
variation
variation
Fig. 19. The problem of incipient species. a: speciation by splitting (allopatric speciation by vicariance). b:
speciation by budding (peripatric speciation). c: aborted speciation. See text 8 6.2.
19
APEX 13(1-2): 1-61, 20 avr. 1998
The recent geological history of the Caribbean
region is one of many important variations of the sea-
level, resulting in huge modifications of the coastlines.
The Western Atlantic and the Eastern Pacific regions
were repeatedly linked by seaways, then isolated again
by emergences of the Panama Isthmus. Many
Caribbean islands were also repeatedly separated, then
reunited, sometimes into new combinations [see maps
figs. 3 (p. 61), fig. 5 (p. 65), fig. 11 (p. 109), fig. 12 (p.
111), fig. 13 (p. 113), fig. 14 (p. 125) in PETUCH 1988;
also maps 2.9 and 2.10 in COATES & OBANDO, 1996)].
These geological circumstances provided an ideal
stage for a scenario of repeated isolations of
populations, reflected in morphological changes (due to
genetic drift or/and ecological factors). These episodes
of isolation were followed by reunions (and aborted
speciations and introgressions). This history also
allowed geographical heterochrony -the persistence of
Neogene faunas in "relict pockets" into the Recent
fauna, [PETUCH 1982, PETUCH 1988: 149-200, map fig.
19 (p. 168)]. In response to such a dynamic geography,
Oliva species in which succesful, long-distance larval
transport and settlement of larva is of weak efficiency
(see above, $ 6.1), can become highly dynamic in their
morphological evolution. It is known that changes can
be rapid: one example has been reported of a small
isolate of an Oliva species [O. amethystina (Rôding,
1798)] undergoing detectable morphological
modifications within the time span of a few decades
(TURSCH 1994).
6.3. Nomenclature: fixed names for dynamic
species.
It could be argued that populations between which the
gene flow is restricted are incipient species and should
therefore be named. It is of course conceivable that
some pairs of such populations, if brought into contact,
could not interbreed anymore but we have yet no
factual evidence to that point.
An apparent paradox is that, when the speciation
issue is undecided, the nomenclatural situation is not.
As said by DAWKINS (1996: 96): "/n a way, our
naming procedures are set up for a pre-evolutionary
age when divides were everything and we did not
expect to find intermediates”. In any case, the Code of
Nomenclature simply has no provision for
hypothetical, future species. If one wishes to draw
attention to such cases, the best one can do is using
infrasubspecific (form) names, which have no
nomenclatural standing.
7. SYSTEMATICS
In this section, the names of very frequently cited
authors will be abbreviated, to save on space. So, B. &
B. stands for BURCH & BURCH, P. & S. for PETUCH &
SARGENT, W. & A. for WAGNER & ABBOTT, Z. & P.
for ZEIGLER & PORRECA. To avoid confusion, “PI.” and
“fig(s).” refer to plates and figures in cited works,
while “PL.” and “FIG(s).” refer to illustrations in the
20
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
present paper. The abbreviation “g.v.” (quod vide)
means “see under that name”.
Throughout this text, the convenient device of a
semicolon inserted between the specific name and the
author [X-us albus ; Smith (not Brown)] is used to
distinguish between a misidentification, which has no
nomenclatural status, and a homonym [X-us albus
Smith (not Brown)], which has (see MAYR & ASHLOCK
1991: 362).
The taxonomical status of every name has been
discussed separately. This entails much repetition but
allows the reader to inquire about a particular taxon
without having to read all the text.
In deference to conchological tradition, the
following "Description" paragraphs are largely based
upon shell ornamentation, which is notoriously variable
in the genus Oliva. So, these sections can serve only for
quick identification. The species have not been
delimited on these characters but on morphometrics
(see $ 4 and 5). This exploratory task being
accomplished, detailed measurements are generally not
indispensable any more for identification of individual
specimens. All characters common to the genus (shell
smooth, shiny, etc.) have been omitted and only
features with some discrimination power are reported.
Preference has been given to details possessing
probably very little or no adaptative value. As just
stated here above, very few (if any) of these traits,
taken one by one, will allow secure identification of an
individual specimen. The probability for achieving this
will be much increased by observing the simultaneous
presence of such features. Oliva species, although
highly variable, have nevertheless limited
“répertoires”. Familiarity with the limits of
intraspecific variation should allow rapid identification
of most (but not all) specimens.
Family OLIVIDAE Latreille, 1825
Subfamily OLIVINAE Latreille, 1825
Genus Oliva Bruguière, 1789
Oliva foxi Stingley, 1984.
Oliva foxi Stingley, 1984: 28.
Description.
SIZE: up to about 40 mm.
SHAPE: fusiform-elongated.
SPIRE: conical, elevated, with large spire callus,
uniformly purple-gray to beige, covering 7 to 4/5
of the whorl.
CHANNEL: rather narrow.
SUBCHANNEL PATTERN: fasciculated, of short brown
stokes.
SHELL BACKGROUND: pale cream to whitish.
COLOUR PATTERN: Faint pink tent pattern overlaid
with distinct dark brown triangular areas, speckled
TURSCH, GREIFENEDER & HUART Oliva fulgurator and related species APEX 13(1-2): 1-61, 20 avr. 1998
with white tents, and heavy brown lines, formed of
rows of small isoscele triangles.
COLUMELLA: white, with faint purple base.
SUPRAFASCIOLAR BAND: lower half with chevron-like
brown marks.
APERTURE: suffused with orange-saffron. Inner
margin of lip milky white.
PROTOCONCH: pink to bright purple-pink.
Diagnosis. Distinguished from all other American
Oliva by its unmistakable, outstanding colour pattern,
saffron aperture and small size.
Distribution. Known only from 18 m, white sand,
Cocos Islands, Costa Rica.
Oliva foxi Stingley, 1984. This taxon, broadly
sympatric with ©. spicata deynzerae is easily
separated from all members of the "O. spicata
complex" (see fig. 3) and is an objective species (see
$ 4.1), in agreement with the conclusions of P. & S.
(1986). No synonyms. The holotype (see PL. 4, FIG.
1) has been previously illustrated by KAICHER 1988
(Part IIL card no. 5249). Three paratypes are
illustrated (see PL. 4, FIGS. 2-4). This species has
been placed by PETUCH & SARGENT (1986: 123) in
their "splendidula group", the argument being: "Both
shells exhibit a very hig gloss and colour patterns that
include bright pinks and purples and wide bands of
dark brown. Both species also have yellow or orange
apertures".
Oliva fulgurator (Rüding, 1798).
Porphyria fulgurator Rôding, 1798: 36, sp. no. 453.
Oliva ispida (Link) (not Rôding, 1798), 1807: 96:
Oliva fusiformis Lamarck, 1811: 318, sp. no. 30;
Encycl. PI 367, figs. la, 1b.
Oliva reticularis Lamarck, 1811: 314, sp. no. 16.
Encycl. PI. 361, figs. 1a,b.
Oliva olorinella Duclos, 1835: PI. 6, figs. 15-16;
text in //lustr. Conch.: 14 (pars).
Oliva obesina Duclos, 1840: PL. 16, figs. 9-11, text
in //lustr. Conch.: 26.
Oliva timoria Duclos, 1840: PI. 17, figs. 11-13;
Illustr. Conch.: 19 (pars).
Oliva aldinia Duclos, 1845: 25, PI .26, figs. 6-7.
Oliva broderipi Ducros de St. Germain, 1857: 62,
sp10:39,; P1/2739;2b:
Oliva jamaicensis Marrat, 1867: 213-15.
Oliva oblonga Marrat, 1867: 215.
Oliva pallida Marrat, 1867: 215.
Oliva bewleyi Marrat, 1870: 7, sp. no. 25, fig. 44.
Oliva figura Marrat, 1870: sp. no. 26, fig. 45.
Oliva formosa Marrat, 1870: sp. no. 16, figs. 29, 30.
Oliva graphica Marrat, 1870: sp. no. 18, PI. 3, fig.
36.
“Oliva oblongata Marrat, 1870” (auct).
Oliva porcea Marrat, 1870: 6, sp. no. 19, PI. 3, fig.
35:
Oliva olivacea Marrat, 1870: 7, sp. no. 28, PI. 4,
figs. 46-47, 51-53.
Oliva hepatica ; Marrat (not Lamarck, 1811), 1871:
sp. no. 14, PI 3, figs. 27-28.
Oliva bullata Marrat, 1871: 40, sp. no..215, PI. 24,
fig. 448.
Oliva circinata Marrat, 1871: sp. no. 109, PI. 17,
fig. 277.
Oliva mercatoria Marrat, 1871: sp. no. 111, PL. 17,
figs. 268-269.
Oliva nivosa Marrat, 1871: sp. no. 112, P1.17, fig.
276; PI. 25, fig. 472.
Oliva reclusa Marrat, 1871: sp. no. 27, PL. 17, fig.
264.
Oliva bifasciata Küster in Weinkauff, 1878: 38, sp.
no. 35, PL. 5, fig. 11; PL 10, figs. 10, 11.
Oliva cribraria Marrat, 1883?. Published in … ?
Oliva reticularis bollingi Clench, 1934: 142, PI. 7,
figs. 3, 4.
Oliva reticularis greenwayae Clench, 1937: 17-26.
Oliva pattersoni, Clench, 1945 4: 49.
Oliva drangai Schwengel, 1951: 117, PL. 8, figs. 2-
3).
Oliva antillensis Petuch & Sargent, 1986: 124, PI.
20, figs. 11, 12.
Oliva bahamasensis Petuch & Sargent, 1986: 125,
pl. 20, figs. 15-18.
Oliva barbadensis Petuch & Sargent, 1986: 126, pl.
20, figs. 19-22.
Oliva bifasciata jenseni Petuch & Sargent, 1986:
128, PI. 21, figs. 16, 17.
Oliva finlayi Petuch & Sargent, 1986: 129, PI 22,
figs. 5-7.
Oliva goajira Petuch & Sargent, 1986: 133, PI. 23,
os 12/13
Oliva jamaicensis zombia Petuch & Sargent, 1986:
136, PL. 24, figs. 17, 18.
Oliva magdae Petuch & Sargent, 1986: 138, PI. 25,
figs. 1-3.
Oliva bifasciata sunderlandi Petuch, 1987: 28, PI.
3, figs. 13, 14.
Oliva circinata tostesi Petuch, 1987: 141.
Oliva sargenti Petuch, 1987: 105, PI. 17, figs. 2, 3.
Oliva maya Petuch & Sargent, 1986: 139, PI. 25,
figs. 4, 5, 8, 9.
Oliva contoyensis Petuch, 1988: 54, PI. 32, figs. 12,
13.
Oliva ernesti Petuch 1990: 63, figs. 19, 20.
Description. Within local populations the shells are
quite homogeneous. The ranges of variations here
under refer to differences between populations.
SIZE: from about 25 mm to over 70 mm.
SHAPE: very variable, from elongated-fusiform to
nearly cylindrical, to globose.
SPIRE: conical to telescopic. Relative height very
variable. Spire callus covering from one half to
nearly all of whorl; shape from flat to convex;
colour from white, beige, purplish to brown, often
darker at upper margin. Callus without distinct
colour strokes.
21
APEX 13(1-2): 1-61, 20 avr. 1998
CHANNEL: of medium width.
SUBCHANNEL PATTERN: fasciculated, very variable.
Not seen in albinistic, melanistic and in rare,
unicoloured specimens.
SHELL BACKGROUND: from white to cream, to yellow,
(O gray.
COLOUR PATTERN: tents and chevrons of very
variable size, width and intensity, sometimes
arranged into vertical series. Colour of markings
varies from pink to reddish, brown or black. In
some populations, markings are diffuse in the
direction of growth. The angle of chevrons is large
and does not vary appreciably with growth (see
Plate 3, fig. 2). Two large spiral bands are formed
by local reinforcement of the pattern. Overlay of
additional uniform brown colour may cover the
whole shell (the so-called "pattersoni pattern", see
PL. 8, FIG. 6), or may appear in two or more sharply
delimited spiral bands (the so-called "bhifasciata
pattern", occurring in widely separated populations,
from Colombia to Florida).
COLUMELLA: from nearly smooth, overlaid with
thick enamel, to heavily plaited. Colour from white
to grevyish, to pale purple.
FASCIOLE: mostly uniform white to greyish, base can
be suffused with colour. Ridges brown in some
populations.
SUPRAFASCIOLAR BAND: very variable.
APERTURE: from white to beige, to faint lilac and
greyish blue. Inner margin of lip: variable, from
aperture colour to row of darker markings. Aspect
depends much on age.
PROTOCONCH: large to extremely large.
Diagnosis. The large values of the protoconch
character RESS and the small NW (number of
nuclear volutions) (see Fig. 8, 1-8) set ©. fulgurator
apart from all other Oliva species, excepted O.
spicata. Distinguished from sayana by constant angle
of chevrons. Distinguished from ©. scripta by a much
narrower channel.
Distribution. Very widely distributed in the Western
Atlantic, from Bermudas to Southern Brazil.
SYNONYMY.
O. fulgurator (Rôding, 1798), based upon the
unambiguous figure 562 of MARTINI (see PL. 6, FIG.
10), is the valid name (see TURSCH, DUCHAMPS &
GREIFENEDER 1994) for an objective species (see $
5.1, 5.4). This is agreement with the conclusions of Z.
& P. (1969), W. & A. (1978) and P. & S. (1986).
This species is very close (or identical) to the fossil
O. schepmani Weissbord, 1962 (not treated here).
The following names are synonyms or designate
local forms:
Oliva ispida (Link) (not Rôding, 1798), 1807. This
taxon (being based upon the same figure 562 of
MARTINI) is an objective junior synonym of O.
fulgurator (Rôding, 1798). It was ©. spicata
(Rôding, 1798) for B. & B. (1960), ©. fulgurator
(Rôding, 1798) for Z. & P. (1969).
22
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
Oliva fusiformis Lamarck, 1811. For original
illustration, see PL. 6, FIG. 5. This has been
demonstrated (see GREIFENEDER, DUCHAMPS &
TURSCH, 1995) to be an objective junior synonym
of ©. fulgurator (Rôding, 1798), in agreement with
B. & B. (1960), Z .& P. (1969), ABBOTT (1974), W.
& A. (1978) and P. & S. (1986). It was ©. reticularis
Lamarck, 1811 for DUCROS de St. GERMAIN (1857).
Oliva reticularis Lamarck, 1811. For the availability of
this name, see GREIFENEDER, DUCHAMPS & TURSCH
(1995). No type material could be located. The
original description 1s:
"16. Olive réticulaire. Oliva reticularis.
O. Cylindracea, alba, subbifasciata; lineis fulvis,
subpunctatis flexuoso-angulatis; spirâ acutä.
Mus., n.12. Encycl., PI. 361, f. 1.
Martini, Conch. 2, t. 51, f 561.
Habite... Sur un fond blanc, cette olive offre quantité
de lignes en zigzags, rousses, subponctuées. Dans les
espaces qu'embrassent deux bandes transverses, ces
lignes, plus épaissies et plus colorées, imitent en
quelque sorte des caractères d'écriture. Cette olive est
peu bombée, a une spire pointue, et ne présente
qu'une couleur blanche à son ouverture. Sa longueur
est d'environ 45 millimètres. Le bord supérieur du
dernier tour est comme dentelé par des taches d'un
brun violet, composé de lignes repliées en faisceau."
The illustration in the Encyclopédie (PI. 361, fig.
1, a, b) depicts a specimen inclined at an angle, as
evidenced by the aspect of the shoulder, the aperture,
and the lower part of the columella. Several other
Oliva in the Encyclopédie are represented in a similar
perspective, which, of course, makes a shell appear
more globose. So the illustration (see PL. 8, FIG. 9)
represents a rather elongated shell, with a peculiar,
very concave lip. The large size (45 mm) and the
straight sides ("peu bombée") indicates that the shell
is not the ©. reticularis of recent authors ("ovate with
rounded sides" in P. & S. 1986).
A shell reasonably matching the original
illustration has been found in the Récluz collection, at
the MHNG, and could possibly be the figured
specimen. Shells of this general type are not
uncommon in some central Caribbean localities.
These forms are linked to the “typical” ©. fulgurator
by an unbroken chain of many intergrading
populations. Considerable efforts invested in
attempting objective separation, by many different
methods and over many years, have always been
completely unsuccessful. Much to our regret (for the
stability of nomenclature) we are compelled to
consider ©. reticularis Lamarck, 1811 as a local form
and a subjective junior synonym of ©. fulgurator
(Rôding, 1798) (see $ 5.1). It was a valid species for
Z. & P. (1969), W. & A. (1978) P. & S. (1986) and
many other authors.
Oliva olorinella Duclos, 1835. The heterogeneous lot
of 8 dirty-white syntypes at MNHN contains
specimens of ©. oliva (L., 1758) (as correctly
inferred by KAICHER 1989, who illustrated one such
syntype on her card Part IV, no. 5516) as well as
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
shells of Caribbean origin (see PL. 9, FIG. 2). These
could not be separated from the “//gurator-
reticularis complex” morphological continuum,
being linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, Oliva olorinella Duclos,
1835 (pars) is here considered as yet another local
variety of O. fulgurator (Rôding, 1798). This was O.
reticularis Lamarck, 1811 for B. & B. (1960), Z. & P.
(1969), ABBOTT (1974), W. & A. (1978) and P. &S.
(1986). This name is often utilised to designate a
whitish, small form from the Bahamas.
Oliva obesina Duclos, 1840. The type material (figured
syntype, out of three, see PL. 6, FIG. 1) could not be
separated from the “/w/gurator-reticularis compiex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of ©.
Julgurator (Rôding, 1798). It is very close to O.
aldinia Duclos, 1845, and to the semi-fossil ©.
schepmani Weissbord, 1962. Three further specimens
have been found in the Duclos collection, at
Clermont-Ferrand, under the unpublished name “O.
opisia”. ©. obesina Duclos, 1840 was ©. spicata
Rôding for B. & B. (1960), Z. & P. (1969), KEEN
(1971), ABBOTT (1974) and W. & A. (1978), a valid
species for P. & S. (1986). Shells very similar to the
type material are found in Venezuela, in the province
of Nueva Esparta.
Oliva timoria Duclos, 1840. The type material is
heterogeneous. One of the syntypes (see PL. 7, FIG. 1)
is very similar to a brown ©. obesina Duclos, 1840
(g.v.), so ©. timoria Duclos, 1840 is (pars) a
subjective junior synonym of ©. fulgurator
(Rôding, 1798), in agreement with Ducros, 1857 (as
©. reticularis Lamarck, 1811).
Oliva aldinia Duclos, 1845. The type series (see PL. 6,
FIG. 2) could not be separated from the “/w/gurator-
reticularis complex” morphological continuum,
being linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, this is here considered as
yet another local variety of ©. fulgurator (Rüding,
1798). This is in agreement with B. & B. (1960), W.
& A. (1978, as "aldina", misspelling) and P. & S.
(1986). Very similar specimens are found in
Venezuela, in the province of Nueva Esparta.
Oliva broderipi Ducros de St. Germain, 1857. The
holotype (illustrated by KAICHER 1989, PART IV, card
no. 5560, and rightly said not to be ©. oliva) is very
badly worn (see PL. 7, FIG. 3) but nevertheless
recognisable shell. It could not be separated from the
“fulgurator-reticularis complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. In the
absence of valid arguments to the contrary, this 1s
here considered as yet another local variety of ©.
Julgurator (Rôding, 1798). It is very close to (if not
identical with) ©. (Strephona) contoyensis Petuch,
1988 (q.v.). ©. broderipi is O. ispidula for B. & B.
(1960), a colour form of ©. oliva taeniata Link for P.
& S. (1986), a form of ©. oliva (L., 1758) for Z. & .P
(1969) and W. & A.. (1978). O. broderipi ; Petuch &
Sargent (not Duclos,1857), 1986 (p. 108, PI. 18, figs.
1, 2) bears no resemblance with the type material and
is an ©. oliva (L., 1758), probably from the Indian
Ocean.
Oliva jamaicensis Marrat, 1867. This taxon was re-
described in the Thesaurus: sp. no. 17, PI. 4, fig. 26.
The holotype (illustrated by KAICHER 1988, Part III,
card no. 5187) (see PL. 6, FIG. 6) resembles ©.
bewleyi Marrat, 1870 and could not be separated
from the “/ulgurator-reticularis complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of ©.
fulgurator (Rôüding, 1798). This was ©. reticularis
Lamarck, 1811 for B. & B. (1960), ©. scripta
Lamarck, 1811 for ABBOTT (1974), possibly ©.
scripta Lamarck, 1811 for W. & A. (1978) and a
valid species for P. & S. (1986).
Oliva oblonga Marrat, 1867. This taxon was re-
described in 1870 in the Thesaurus (sp. no. 11, PI. 2,
fig. 14). The large holotype (illustrated by KAICHER
1989, Part IV, card no. 5484) (see PL. 6, FIG. 7) could
not be separated from the “/w/gurator-reticularis
complex” morphological continuum, being linked to
all other conspecific forms by many intergrading
populations. In the absence of valid arguments to the
contrary, this is here considered as yet another local
variety of ©. fulgurator (Rôding, 1798). This was ©.
spicata (Rôding, 1798) for B. & B. (1960), KEEN
(1971) and W. & A. (1978); a colour form of ©.
tisiphona Duclos, 1845 for P. & S. (1986) and
PETUCH (1987), ©. bewleyi Marrat, 1870 for DIAz &
PUY ANA (1994).
Oliva pallida Marrat, 1867. The taxon was re-described
in the Zhesaurus: 27, sp. no. 138, PL. 21, figs. 341-
343. The type material is missing, as already reported
by MCMILLAN (1985). But the identity of the taxon
leaves little doubt because Marrat himself considers it
as a synonym of his own ©. nivosa (q.v.). His label
for the type material of ©. nivosa Marrat, 1871 reads:
“O. nivosa 112 Marrat pallida Marrat in Annals &
Mag. of N. History”. In the Index of the Thesaurus
one also reads: “pallida Marr., nivosa, Marr.” It
follows that Oliva pallida Marrat, 1867 is a local
variety of O. fulgurator (Rôüding, 1798), linked to all
other conspecific forms by many intergrading
populations. This was ©. reticularis Lamarck, 1811
for B. & B. (1960), Z. & P. (1969), ABBOTT (1974)
and P. & S. (1986), ©. olorinella Duclos, 1835 for
W. & A. (1978). Note: the name pal/lida has been
also used by SWAINSON (1831: 78, PL 3, fig. 2) for a
species of Agaronia, by DAUTZENBERG (1910) for a
variety Of Oliva flammulata Lamarck, 1811 and by
DAUTZENBERG (1927: 71: 110) for a colour form of
Oliva reticulata (Rôding, 1798).
Oliva bewleyi Marrat, 1870. The holotype (illustrated
by KAICHER 1989, Part IV, card no. 5557) (see PL. 7,
23
APEX 13(1-2): 1-61, 20 avr. 1998
FIG. 10) could not be separated from the “/ulgurator-
reticularis complex” morphological continuum,
being linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, this is here considered as
yet another local variety of ©. fulgurator (Rüding,
1798). This is ©. reticularis Lamarck, 1811 for B. &
B. (1960) and W. & A. (1978), a valid species for P.
& S. (1986) and Diaz & PUYANA (1994) who
insisted on the extreme variation of this taxon.
Oliva figura Marrat, 1870. The holotype (see PL. 8,
FIG. 3) (illustrated by KAICHER 1988, Part III, card
no. 5186) could not be separated from the
“fulgurator-reticularis complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. In the
absence of valid arguments to the contrary, this is
here considered as yet another local variety of ©.
Julgurator (Rôding, 1798). This was ©. reticularis
Lamarck, 1811 for B. & B. (1960) and W. & A.
(1978). It was a colour form of “O. jamaicensis
brunnea Marrat” for P. & S. (1986).
Oliva formosa Marrat, 1870. The slim holotype, with
several brown spiral stripes (illustrated by KAICHER
1989, Part IV, card no. 5566) (see PL. 9, FIG. 6),
could not be separated from the “/w/gurator-
reticularis complex” morphological continuum,
being linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, this is here considered as
yet another local variety of ©. fulgurator (Rôüding,
1798). This was ©. reticularis Lamarck, 1811 for B.
& B. (1960) and Z. & P. (1969), a form of the same
for ABBOTT (1974), W. & A. (1978) and a colour
form of ©. bewleyi Marrat, 1870 for P & S (1986).
Oliva graphica Marrat, 1870. The swollen, worn
holotype (see PL. 6, FIG. 4) (illustrated by KAICHER
1989, Part IV, card no. 5561) could not be separated
from the “fulgurator-reticularis complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of ©.
fulgurator (Rôding, 1798). This was ©. obesina
Duclos, 1840 for TOMLIN (in FORD 1953), ©. spicata
(Rôding, 1798) for B. & B. (1960), ©. scripta
Lamarck, 1811 for OLD (cited in Z & P, 1969),
Abbott (1974) and W. & A. (1978); ©. julieta
Duclos, 1840 for Z. & P. (1969) and Keen (1971). It
was à valid species for P. & S. (1986: 134, PI. 23,
figs. 7, 8) but their illustration does not at all match
the type material.
“Oliva oblongata Marrat, 1870” (auct). This is a
misspelling (and a date error) for ©. oblonga Marrat,
1867 (q.v.). It was ©. spicata (Rôding, 1798) for Z. &
P. (1969), W. & A. (1978), Abbott (1974) and P. & S.
(1986).
Oliva olivacea Marrat, 1870. This name is available, as
O. olivaceus Meuschen, 1787 is a nomen nudum (in a
rejected work) as noted by W. & A. (1978). The same
24
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
would apply for ©. olivacea Karsten, 1789, should
the work of that author be rejected by the
Commission. Marrat’s taxon (illustrated by KAICHER
1988, Part III, card no. 5163), reported as Oliva
olivacea Meuschen, is certainly a local variety of ©.
Julgurator (Rôding, 1798), linked to all other
conspecific forms by many intergrading populations.
It was indeed ©. reticularis Lamarck, 1811 for B. &
B. (1960), Z. & P. (1969), ABBOTT (1974), W. & A.
(1978), ©. bewleyi Marrat, 1870 for DIAZ & PUY ANA
(1994). It was a valid species for P & S (1986).
Marrat himself wrote ‘“olivaceus Meuschen:;
reticularis Lamk.” in his caption to Plate IV.
Oliva porcea Marrat, 1870. The very fat holotype (see
PL. 6, FIG. 3) is quite similar to ©. obesina Duclos,
1840, as already stated by TOMLIN (in FORD 1953). It
could not be separated from the “/w/gurator-
reticularis complex” morphological continuum,
being linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, this is here considered as
yet another local variety of ©. fulgurator (Rüding,
1798). This was ©. spicata (Rôding, 1798) for B. &
B. (1960), ©. scripta Lamarck, 1811 for ABBOTT
(1974) and W. & A. (1978); ©. julieta Duclos, 1840
for Z. & P. (1969) and KEEN (1971).
Oliva hepatica ; Marrat (not Lamarck, 1811), 1871.
The shell described in Marrat as ©. hepatica Lamarck
does not correspond to the original, very vague
description of Lamarck. Marrat’s figures leave little
doubt that this is the same as ©. bifasciata Küster
1878, as already suggested by Z. & P. (1969) and W.
& A. (1978). This was ©. reticularis Lamarck, 1811
for B. & B. (1960), possibly a form of the same for Z.
& P. (1969), a colour form of ©. tisiphona Duclos,
1845 for P. & S. (1986). It is in any case a local
variety of O. fulgurator (Rôding, 1798). O. hepatica
Lamarck, 1811 is a nomen dubium (see
GREIFENEDER, DUCHAMPS & TURSCH 1995).
Oliva bullata Marrat, 1871. The holotype (1llustrated
by KAICHER 1988, Part III, card no. 5172) (see PL. 9,
FIG. 3) is a local variety of ©. fulgurator (Rôding,
1798) could not be separated from the “/w/gurator-
reticularis complex” morphological continuum,
being linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, this is here considered as
yet another local variety of ©. fulgurator (Rôding,
1798). This was ©. fulgurator (Rüding, 1798) for B.
& B. (1960) and W. & A. (1978), a subspecies of the
same for P. & S. (1986).
Oliva circinata Marrat, 1871. The holotype (see PL. 9,
FIG. 8) (illustrated by KAICHER 1988, Part III, card
no. 5526), the four paratypes at MCM, and a probable
paratype at AMNH form an homogeneous series that
could not be separated from the “/w/gurator-
reticularis complex” morphological continuum,
being linked to all other conspecific forms by many
intergrading populations. This rather distinctive form
very often occupies a peripheral position in scatter
diagrams. It also occupies a large, peripheral
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
geographic range all along the coast of Brazil, South
of the possible geographical barrier formed by the
Amazon River. All populations in this region share
the same characteristics, with very little variation
save that some (for instance one found in 30 m off
Recife) are smaller in size. The temptation to
consider this as a subspecies was resisted because
extremely similar forms occur in deep water off
Florida, in Venezuela and in Colombia. So this taxon
is here considered to be a local variety of ©.
Julgurator (Rôding, 1798). O. circinata Marrat, 1871
was ©. sayana Ravenel, 1834 for B. & B. (1960), Z.
& P. (1969), ABBOTT (1974) and W. & A. (1978); a
colour form of ©. graphica Marrat, 1870 for P. & S.
(1986), O. figura Marrat, 1870 for DIAZ & PUY ANA
(1994). ©. circinata Martyn, 1789, a nomen nudum
(in a rejected work) was ©. lignaria Marrat, 1868 for
W. & A. (1978).
Oliva mercatoria Marrat, 1871. The two syntypes at
MCM (one figured PL. 6, FIG. 9) could not be
separated from the “/u/gurator-reticularis complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of ©.
Jfulgurator (Rôding, 1798). This was also the
interpretation of B. & B. (1960), Z. & .P (1969), W.
& A. (1978) and P. & S. (1986).
Oliva nivosa Marrat, 1871. The holotype at MCM
(illustrated by KAICHER 1988, Part III, cards no.
5158, 5159) (see PL. 7, FIG. 9) and one possible
paratype could not be separated from the “fu/gurator-
reticularis complex” morphological continuum,
being linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, this is here considered as
yet another local variety of ©. fulgurator (Rüding,
1798). This was ©. reticularis Lamarck, 1811 for B.
& B. (1960), Z. & .P (1969), ABBOTT (1974) and W.
& A. (1978), a valid species for P. & S. (1986).
Oliva reclusa Marrat, 1871. The holotype (see PL. 9,
FIG. 7), illustrated as “syntype” by KAICHER 1988,
Part III, (card no. 5247) was correctly described by
TOMLIN (in FORD 1953) as “a pale reticularis Lam. It
could not be separated from the “/w/gurator-
reticularis complex” morphological continuum,
being linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, this is here considered as
yet another local variety of ©. fulgurator (Rôüding,
1798). Very similar specimens are found in Aruba
(see HEMMEN 1981). This was ©. reticularis
Lamarck, 1811 for B. & B. (1960), Z. & P. (1969),
ABBOTT (1974) and W. & A. (1978), a valid species
for P. & S. (1986) and PETUCH (1987).
Oliva bifasciata Küster in Weinkauff, 1878. The
locality given is “South coast of America (Marrat),
probably Brazil and Guyana”. The reference "PI. 10,
figs. 8,9" in the text is wrong and is corrected to " PI.
10, Figs. 10, 11" in the "Erklärung der Tafeln". The
figured specimen (H: 61.39 mm, D: 26.73 mm) (see
PL. 8, FIG. 4) SMF no. 9353 represents a striking, but
common colour variant, encountered in many
Caribbean populations. It could not be separated from
the “fulgurator-reticularis complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. In the
absence of valid arguments to the contrary, this is
here considered as a colour form of ©. fulgurator
(Rôding, 1798). ©. bifasciata Küster in Weinkauff,
1878 was ©. reticularis Lamarck, 1811 for B. & B.
(1960), Z. & P. (1969), ABBOTT (1974), W. & A.
(1978), a separate, valid species for P. & S. (1986).
Oliva cribraria Marrat, 1883? This taxon is
represented by a properly labelled holotype in MCM.
The work in which the publication appeared is
unknown. According to MCMILLAN (1985)
"Description and col. fig. of this species exist in print
(two copies), possibly ex Marrat's privately printed
Notebook of a Liverpool Naturalist (1833)". The
upper edge of the fasciole of the beautiful holotype
(see PL. 9, FIG. 10) is delimited by a thin, bright
purple zone (more vivid than in ©. circinata Marrat,
1871, to which the shell is very closely related). The
type (with a purple protoconch) could not be
separated from the “fulgurator-reticularis complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of ©.
Julgurator (Rôding, 1798). Locality unknown. This
was ©. oblonga Marrat for TOMLIN in FORD (1953).
Oliva reticularis bollingi Clench, 1934. The heavy
holotype (see PL. 7, FIG. 8) (illustrated by KAICHER
1989, Part IV, card no. 5543) could not be separated
from the “fulgurator-reticularis complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of ©.
Julgurator (Rôding, 1798). ©. bollingi Clench, 1934
was ©. reticularis Lamarck, 1811 for B. & B. (1960),
Z. & P. (1969), ABBOTT (1974) and W. & A. (1978);
a subspecies of ©. bifasciata Küster, 1878 for P. &S.
(1986). This form comes from crab traps, off Miami,
Florida.
Oliva reticularis greenwayae Clench, 1937. The
holotype (see PL. 8, FIG. 5) (illustrated by KAICHER
1989, Part IV, card no. 5562) is extremely similar to
the figured specimen of ©. bifasciata Küster in
Weinkauff, 1878 (see PL. 8, FIG. 4). It could not be
separated from the “/u/gurator-reticularis complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety (and
colour form) of ©. fulgurator (Rôding, 1798). ©.
reticularis greenwayae Clench, 1937 was ©.
reticularis Lamarck, 1811 for B. & B. (1960), Z. & P.
(1969) and ABBOTT (1974); ©. bifasciata Küster,
25
APEX 13(1-2): 1-61, 20 avr. 1998
1878 for W & A (1978) and P. & S. (1986). This
form comes from Smith Point, Grand Bahama Island,
Bahamas.
Oliva reticularis pattersoni, Clench, 1945. The dark,
heavy holotype (see PL. 8, FIG. 6) (illustrated by
KAICHER 1989, Part IV, card no. 5568) could not be
separated from the “/u/gurator-reticularis complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety (and
colour form) of ©. fulgurator (Rôüding, 1798). O.
pattersoni Clench, 1945 was ©. reticularis Lamarck,
1811 for B. & B. (1960), Z. & P. (1969) and ABBOTT
(1974), ©. formosa Marrat, 1870 for W. & A. (1978)
and KAICHER (1989), a colour form of ©. bifasciata
bollingi Clench, 1937 (sic) for P. & S. (1986). This
form comes from 5 miles E of Crabbing Point, Great
Bahama Island, Bahamas.
Oliva drangai Schwengel, 1951. The light-coloured
holotype (see PL. 7, FIG. 7) (previously illustrated by
KAICHER 1988, Part IIL card no. 5165, as “O.
drangae”) could not be separated from the
“fulgurator-reticularis complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. In the
absence of valid arguments to the contrary, this 1s
here considered as yet another local variety of ©.
fulgurator (Rôding, 1798). ©. drangai Schwengel,
1951 was probably ©. scripta Lamarck, 1811 for W.
& A .(1978), a valid species for P. & S. (1986). This
form comes from Pigeon Pt, Tobago. Note: the
holotype (ANSP 247107) has an abnormal, tilted
protoconch. The protoconch of the smaller (H: 18.56
mm, D: 8.46 mm) paratype (ANSP 247093) is
normal.
Oliva antillensis Petuch & Sargent, 1986. The holotype
(see PL. 9, FIG. 1) (illustrated by KAICHER 1988, Part
III, card no. 5245) could not be separated from the
“fulgurator-reticularis complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. In the
absence of valid arguments to the contrary, this 1s
here considered as yet another local variety of O.
fulgurator (Rôding, 1798). This form comes from the
Southern coast of Gonave Island, HAITI. This was ©.
bewleyi Marrat for DIAZ & PUY ANA (1994).
Oliva bahamasensis Petuch & Sargent, 1986. The
holotype (see PL. 8, FIG. 10) (illustrated by KAICHER
1989, Part IV, 5575) could not be separated from the
“fulgurator-reticularis complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. In the
absence of valid arguments to the contrary, this is
here considered as yet another local variety of ©.
fulgurator (Rôding, 1798). This form was caught in a
lobster pot, in 200 m off Grand Bahama I.
Oliva barbadensis Petuch & Sargent, 1986. The
holotype (see PL. 9, FIG. 4) (illustrated by KAICHER
1989, Part IV, card no. 5531) could not be separated
26
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
from the “fulgurator-reticularis complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of ©.
Julgurator (Rôding, 1798). This form comes from
deep water. (200 m) off St. James, Barbados. Other
deep water specimens from close neighbouring
localities have a very different colour pattern.
Oliva bifasciata jenseni Petuch & Sargent, 1986. The
holotype (see PL. 7, FIG. 6) (illustrated by KAICHER
1988, Part III, card no. 5202) could not be separated
from the “/ulgurator-reticularis complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of ©.
Julgurator (Rôüding, 1798). This form was collected
in shallow water (0.5 m), in Bermuda.
Oliva finlayi Petuch & Sargent, 1986. The holotype
(see PL. 8, FIG. 8) (illustrated by KAICHER 1988, Part
III, card no. 5184) could not be separated from the
“fulgurator-reticularis complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. In the
absence of valid arguments to the contrary, this is
here considered as yet another local variety of ©.
Jfulgurator (Rôding, 1798). This is a deep water form,
from 200 m, Matanzas Bay, Cuba.
Oliva goajira Petuch & Sargent, 1986. The holotype
(see PL. 8, FIG. 2) (illustrated by KAICHER, 1988, Part
III, card no. 5189) could not be separated from the
“fulgurator-reticularis complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. In the
absence of valid arguments to the contrary, this is
here considered as yet another local variety of O.
fulgurator (Rôüding, 1798). This form comes from 60
m off Cabo La Vela, Colombia. It falls within the
range of variation of ©. circinata Marrat, 1871 for
Diaz & PUYANA (1994). An extremely similar form
lives in 5-6 m, coarse sand, Tayrona, Colombia.
Oliva jamaicensis zombia Petuch & Sargent, 1986.
The holotype (see PL. 7, FIG. 2) could not be
separated from the “/y/gurator-reticularis complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of ©.
fulgurator (Rôüding, 1798). This form was collected
on sand flats, Southern side of Gonave I., Haiti.
Oliva magdae Petuch & Sargent, 1986. The holotype
(see PL. 6, FIG. 8) could not be separated from the
“fulgurator-reticularis complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. In the
absence of valid arguments to the contrary, this is
here considered as yet another local variety of ©.
fulgurator (Rôding, 1798). This form was caught in a
fish trap, 300 m, off Matanzas Bay, Cuba.
TURSCH, GREIFENEDER & HUART
Oliva maya Petuch & Sargent, 1986. The holotype (see
PL. 8, FIG. 1) (illustrated by KAICHER 5186) could not
be separated from the “/w/gurator-reticularis
complex” morphological continuum, being linked to
all other conspecific forms by many intergrading
populations. In the absence of valid arguments to the
contrary, this is here considered as yet another local
variety of O. fulgurator (Rüding, 1798). This form
was trawled by a shrimper in 35 m. off Contoy Is.
Yucatan, Mexico. It is broadly sympatric with Oliva
contoyensis Petuch, 1988, but there is no clear
indication that the two taxa are syntopic. For reasons
unknown to us, ©. maya was considered by PETUCH
& SARGENT (1986: 121) to belong to their "sayana
group".
Oliva bifasciata sunderlandi Petuch, 1987. The
holotype (see PL. 9, FIG. 5) could not be separated
from the “fulgurator-reticularis complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of ©.
Jfulgurator (Rôding, 1798). This form was trawled
from 150 m West of Cedar Key, Florida.
Oliva circinata tostesi Petuch, 1987. The holotype (see
PL. 9, FIG. 9) (illustrated by KAICHER 1989, Part IV,
card no. 5530) and several topotypes could not be
separated from a large sample of ©. fulgurator
circinata Marrat, 1871, encompassing several
populations. This, being described from a single
population, does hardly qualify as a subspecies and is
(at best) a local variety of ©. fulgurator (Rôding,
1798).
Oliva sargenti Petuch, 1987. The holotype (see PL. 7,
FIG. 5) could not be separated from the “fu/gurator-
reticularis complex” morphological continuum,
being linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, this is here considered as
yet another local variety of ©. fulgurator (Rôüding,
1798). This form was collected in 5 m, Malmok,
Aruba.
Oliva contoyensis Petuch, 1988. The holotype (see PL.
7, FIG. 4) is very similar to that of ©. broderipi
Ducros de St. Germain, 1857 (q.v.). It could not be
separated from the “fulgurator-reticularis complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of ©.
Julgurator (Rôding, 1798). This form was collected
in 35 m. off Contoy I., Yucatan, Mexico. It is broadly
sympatric with Oliva maya Petuch & Sargent, 1986,
but there is no clear indication that the two taxa are
syntopic.
Oliva ernesti Petuch 1990. The holotype (see PL. 8,
FIG. 7) could not be separated from the “/u/gurator-
reticularis complex” morphological continuum,
being linked to all other conspecific forms by many
intergrading populations. In the absence of valid
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
arguments to the contrary, this is here considered as
yet another local variety of ©. fulgurator (Rôding,
1798). This form was trawled in 40 m, silty sand, off
Portobello, Panama (East Coast).
The following names have been mistakenly (or very
questionably) considered as synonyms:
Oliva oriola ; Duclos (not Lamarck, 1811), 1835: PI.
10, figs. 1, 2, text in Z/lustr. Conch.: 15. This was ©.
spicata (Rôding, 1798) and ©. reticularis Lamarck,
1811 for W & A (1978). It was ©. spicata (Rôding,
1798) for B. & B. (1960), Z. & P. (1969) and KEEN
(1971). Duclos’ illustrations of ©. oriola Lamarck,
1811 are somewhat ambiguous and, in the absence of
specimens in the Duclos collection, attribution is
most uncertain. It would be better to consider this as
a nomen dubium. In any case, this is a junior
homonym of ©. oriola Lamarck, 1811 (see
GREIFENEDER, DUCHAMPS & TURSCH 1995).
Oliva quersolina Duclos, 1835: PI. 10, figs.7-8. This is
O. olorinella Duclos, 1835 for W. & A. (1978) and
O. reticularis Lamarck, 1811 for Z. & P. (1969),
ABBOTT (1974) and P. & S. (1986). ©. quersolina has
been demonstrated to be a subjective junior
synonym of the Indian Ocean species ©. afalina
Duclos, 1835 (see TURSCH & GREIFENEDER 1996).
Oliva memnonia Duclos, 1845: 15, PI. 17, figs. 19-20.
This was ©. reticularis Lamarck, 1811 for B. & B.
(1960), Z. & P. (1969), ABBOTT (1974) and W. & A
(1978). The original figures (illustrated by KAICHER
1989, Part IV, card no. 5497) are not recognisable
with any certainty. In the absence of type material, it
is much safer to consider this name as a nomen
dubium.
Oliva tisiphona Duclos, 1845. No type material has
been located. The characteristic sub-channel pattern
of the “/ulgurator-reticularis complex” ïs not
apparent on the illustrations of DUCLOS and is not
mentioned in the original description. Although
DUCLOS says this taxon is rather close to ©. oniska
Duclos, 1845, his illustrations are quite problematic
and it is certainly safer to consider this as a nomen
dubium. This was ©. reticularis Lamarck, 1811 for
B. & B. (1960), Z. & P. (1969), ABBOTT (1974) and
W. & A .(1978). P. & S. (1986) consider this to be a
valid species, but it is not at all evident that the
specimen they illustrate (PI. 29, figs. 11-12) matches
the original figure.
"Oliva alba Lamarck" was listed, without a word of
clarification, by GRAY (1858: 44) in the synonymy of
Strephona reticularis. This is a misquotation, as
there is no ©. alba Lamarck (see GREIFENEDER,
DUCHAMPS & TURSCH 1995). Gray refers to sp. no.
42 (O. candida) of Lamarck for which "a/ba" is part
of the Latin description (see B. & B. 1960).
"Oliva vermiculata Lamarck" was listed, without a
word of clarification, by GRAY (1858: 44) in the
synonymy of Sfrephona reticularis. This is a
misquotation, as there is no ©. vermiculata Lamarck
(see GREIFENEDER, DUCHAMPS & TURSCH 1995).
21
APEX 13(1-2): 1-61, 20 avr. 1998 Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
“Oliva diaphana Duclos 1835” (auct) was ©.
reticularis Lam. for B & B (1960), W & A (1978)
and P. & S. (1986). This is a misquotation, as there
is no Oliva diaphana described by Duclos.
"Oliva vermiculata Gray, 1858" (auct.) constitutes a
fine example of sequential taxonomic hallucinations
in the genus Oliva. This non-existent taxon was ©.
reticularis Lamarck, 1811 for B. & B. (1960) and W.
& A. (1978). It was considered to be a valid species
by P. & S. (1986: 155). PETUCH (1987: 105) even
described an Oliva from Haiti as “closely resembling
Gray's type of ©. vermiculata”. One wonders where
this type material could be seen. "Oliva vermiculata
Gray, 1858" is a misquotation because there is no ©.
vermiculata Gray. One finds a citation of
“"vermiculata Lamarck" in Gray (1858: 44), in the
synonymy of Sfrephona reticularis). But this is yet
another misquotation, as there is no such thing as a
“O. vermiculata Lamarck” (q.v.).
Oliva sowerbyi Marrat, 1870: 13, sp. no. 61, PL 8, figs.
114,115. This was ©. reticularis Lamarck, 1811 for
B. & B. (1960), Z. & P. (1969), ABBOTT (1974), W.
& A. (1978) and P. & S. (1986). Oliva sowerbyi
Marrat, 1870 (not illustrated here) has been
demonstrated to be a subjective junior synonym of
the very different Indian Ocean species ©. atalina
Duclos, 1835 (see TURSCH & GREIFENEDER, 1996).
Oliva sowerbyi Anton, 1839: 102 is does not to the
genus Oliva but is a small fossil, possibly an Ancilla.
(see TURSCH & GREIFENEDER, 1996). Likewise.
Oliva sowerbyi Ducros de St-Germain, 1857: 105, sp.
no. 103 is not an Oliva but an Olivella.
Oliva polpasta Duclos, 1833.
Oliva polpasta Duclos, 1833: PL 20; 1840: PL. 16, figs.
1-2: 1844: 26.
Oliva truncata Marrat, 1867: 215.
Oliva kerstitchi da Motta, 1985: 8-9.
Oliva olssoni Petuch & Sargent, 1986: 140, PI. 25,
figs. 17, 18.
Description.
SIZE: to over 50 mm.
SHAPE: ovate to biconic.
SPIRE: flat conical. Spire callus covers from 1/2 to
nearly all the whorl. Colour of callus from bluish
grey to yellow, beige and brown, often darker in
upper zone; occasional dark strokes or points.
CHANNEL: medium.
SUBCHANNEL PATTERN: fasciculated, forming a
regular, narrow zone of black triangles pointing
upwards alternating with whitish triangles of equal
size, pointing downwards. When seen in apical
view, this produces a characteristic “cogwheel
pattern” (see PL. 3, FIG. 6).
SHELL BACKGROUND: yellowish brown to olive gray
to bluish gray.
COLOUR PATTERN: network of chevrons of variable
thickness (mostly blurred to produce nearly
homogeneous backdrop), overlaid with a pattern of
28
single dark blotches, sometimes forming strokes,
chevrons or ziczacs. Two faint large spiral bands
(not always present) are formed by local
reinforcement of the pattern. Some specimens have
one or two wide spiral white bands (“kerstitchi
pattern”).
COLUMELLA: rather smooth, mostly white, with very
faint yellow to green shadow only at the base.
FASCIOLE: uniform white.
SUPRAFASCIOLAR BAND: lower part often with nearly
axial, sharp strokelets.
APERTURE: white to pale greyish or yellowish. Inner
margin of lip: like aperture; dark brown or grey
only in shells with a sharp lip.
PROTOCONCH: medium large.
Diagnosis. Differs from ©. spicata by the all-white
fasciole, the “cogwheel pattern” (see PL. 3, FIG. 6)
and a protoconch generally smaller.
SYNONYMY.
Oliva polpasta Duclos, 1833. This taxon (of which the
figured syntype is illustrated PL. 4, FIG. 6), although
closely related to ©. spicata (Rôding, 1798), is easily
and totally separated from that species (see TURSCH
& HUART, 1990) by quantitative characters. Both are
syntopic (for instance at Cebaco IL, Panama) and
Oliva polpasta Duclos, 1833 thus is the valid name
for a separate, objective species, in agreement with
Z. & P. (1969), W. & A. (1978) and P. & S. (1986). It
was a variety of ©. spicata (Rôüding, 1798) for B. &
B. (1960). This species is very close (or identical) to
the fossils ©. davisae Durham, 1950 and ©. callosa
Li, 1930 (not treated here).
The following names are synonyms or designate
local forms:
Oliva truncata Marrat, 1867. The holotype (see PL. 4,
FIG. 7) could not be separated from ©. polpasta
Duclos, 1833. In the absence of valid arguments to
the contrary, this is here considered as a subjective
junior synonym of ©. polpasta Duclos, 1833. This
was a valid species for P. & S. (1986). It was ©.
elegans Lamarck, 1811 (a completely different Indo-
Pacific species, see GREIFENEDER, DUCHAMPS &
TURSCH 1995) for B. & B. (1960) Z. & P. (1969) and
W. & A. (1978).
Oliva kerstitchi da Motta, 1985. The holotype (see PL.
4, FIG. 8) (with the label "kirstitchi) could not be
separated from ©. polpasta Duclos, 1833, except for
the presence of a white spiral band on the body
whorl. TURSCH & HUART (1990) considered this as
having specific status, as a working hypothesis,
awaiting further confirmation (see $ 1.1.3). Further
work (see $ 4.1) established that ©. kerstitchi da
Motta, 1985 is consistently syntopic with ©. polpasta
Duclos, 1833, to which it is linked by an unbroken
chain of intergrades (KOCH 1992). In the absence of
valid arguments to the contrary, this is here
considered as a colour form of ©. polpasta Duclos,
1833. This was a valid species for P. & S. (1986).
TURSCH, GREIFENEDER & HUART
Oliva olssoni Petuch & Sargent, 1986. The faded,
bulging holotype (see PL. 4, FIG. 9) presents the
characteristic sub-channel "cogwheel pattern" and
could not be separated from ©. polpasta Duclos,
1833, to which it is linked by an unbroken chain of
intergrades (bulging forms of ©. polpasta are not
uncommon, for instance in Panama). In the absence
of valid arguments to the contrary, this is here
considered as a local variety of ©. polpasta Duclos,
1833.
Oliva sayana Ravenel, 1834.
Oliva sayana Ravenel, 1834: 19.
Oliva litterata Lamarck (not Rôding, 1798), 1811:
315, sp. no. 20.
Oliva circinata var. citrina Johnson, 1911: 23.
Oliva (Strephona) sayana sarasotensis Petuch &
Sargent, 1986: 146, PI. 28, figs. 4, 5.
Oliva (Strephona) sayana texana Petuch & Sargent,
1986: 147, PL. 38, figs. 3, 4.
Description.
SIZE: to over 70 mm.
SHAPE: elongated fusiform to cylindrical. In old
spécimens, anterior part of lower lip very heavy and
extended outwards.
SPIRE: conical and distinctly telescopic. Spire callus
covering only 1/3 or 1/2 of the whorl. Colour of
callus beige or gray to orange, with no markings.
CHANNEL: medium.
SUBCHANNEL PATTERN: Coarsely fasciculated, very
rapidly shifting to tent pattern.
SHELL BACKGROUND: whitish-grey to yellow.
COLOUR PATTERN: tents and chevrons of variable
size, sometimes arranged into vertical series. Very
large white tents are common. Two large spiral
bands are formed by local reinforcement of the
pattern. The angle of chevrons starts small and does
appreciably increase with growth (see PL. 3, FIG.
1). Golden forms occur, with overall yellow
appearance and reduction of the pattern.
COLUMELLA: with very strong coarse plaits, often
smoothed by enamel. Colour white.
SUPRAFASCIOLAR BAND: very variable.
APERTURE: inner part often lilac to pink, changing to
whitish grey or cream towards the lip. Inner margin
of lip: dark (interrupted or not) in shells with sharp
lip.
PROTOCONCH: small.
Diagnosis. Differs from ©. fulgurator and ©. scripta
by a marked increase of the angle of chevrons during
growth (see PL. 3, FIGS. 1, 2), a much smaller
protoconch, a telescopic spire. Differs from ©.
scripta by a much narrower channel.
Distribution. Southeastern coast ud U.S. and Gulf of
Mexico. According to P. & S. (1986: 121), ©. sayana
"cannot tolerate the tropical carbonate environment
of the southern tip of Florida … [which] acts as a
barrier to dispersal"
Oliva fulgurator and related species APEX 13(1-2): 1-61, 20 avr. 1998
SYNONYMY.
Oliva sayana Ravenel, 1834. The type is missing in the
Ravenel collection, housed at the Charleston Museum
(ide Dr. Harry D. LEE, in lift). There is no original
figure, nor any actual description. Comparing this
shell to ©. litterata (described by Lamarck as being
66 to 68 mm -about 2.75 inches- long), Ravenel
writes:
“These Shells are certainly distinct, and
therefore should be distinguished by different
names. The ©. sayana, sometimes exceeds 3
inches in length -fine specimens are rare- worn
spécimens are not uncommon on the coast of
South Carolina.”
With so little information, this widely used name
should normally be a nomen dubium. Nomenclatural
stability can nevertheless be preserved by an unusual
(but good) argument: no other Oliva species is found
on the coast of South Carolina (a very intensively
prospected area). This species is easily and
completely separated by quantitative criteria from all
other Atlantic Oliva species (see TURSCH &
HUART,1990), so one can consider that Oliva sayana
Ravenel, 1834 is the valid name for an objective
species, in agreement with Z. & P. (1969), W. & A.
(1978) and P. & S. (1986).
The following names are synonyms or designate
local forms:
Oliva litterata Lamarck, 1811. This (see original
illustration PL. 5, FIG. 1) is the former name of ©.
sayana Ravenel, 1834, in agreement with B. & B.
(1960), Z. & P. (1969), W. & A. (1978) and .P & S.
(1986). It is a junior homonym of ©. lfterata
(Rôüding, 1798) (see TURSCH & al., 1994), a synonym
of ©. spicata (Rôding, 1798) for KEEN (1971), a
nomen dubium for TURSCH, DUCHAMPS &
GREIFENEDER, 1994.
Oliva circinata var. citrina Johnson, 1911.
Examination of the paratype MCZ 6267 (figured here
PL. 5, FIG. 4) (illustrated by KAICHER 1989, Part IV,
card no. 5482) confirms this is a colour form of ©.
sayana Ravenel, 1834, in agreement with B. & B.
(1960), Z. & P. (1969), W. & A. (1978) and P &S.
(1986). This conclusion is supported by the
characteristic small, intact protoconch.
Oliva sayana sarasotensis Petuch & Sargent, 1986.
The rather small (40 mm) holotype (see PL. 5, FIG. 3)
(previously illustrated by KAICHER 1989, Part IV,
card no. 5493) could not be separated from a large
sample of ©. sayana Ravenel, 1834, containing many
populations. This, being described from a single
population (15 km West of Lido Beach, Sarasota,
Florida), does hardly qualify as a subspecies. In the
absence of valid arguments to the contrary, it is here
considered as a local variety of O. sayana Ravenel,
1834.
Oliva sayana texana Petuch & Sargent, 1986. The
holotype (see PL. 5, FIG. 2) (previously illustrated by
KAICHER 1989, Part IV, card no. 5487) could not be
29
APEX 13(1-2): 1-61, 20 avr. 1998
separated from a large sample of ©. sayana Ravenel,
1834, containing many populations. This, being
described from a single population (shallow water,
Padre I., off South Texas), does hardly qualify as a
subspecies. In the absence of valid arguments to the
contrary, it 1s here considered as a local variety of ©.
sayana Ravenel, 1834.
The following names have been mistakenly
considered as synonyms:
Oliva polita Marrat, 1867. The two small, dark
syntypes (with no locality data) at MCM (one
illustrated by Kaicher 1988, Part III, card no. 519)
do not at all resemble ©. sayana Ravenel, 1834 and
are most probably the Polynesian form of ©
panniculata Duclos, 1835, an extremely different,
well-known Indo-Pacific shell. This was ©. sayana
Ravenel, 1834 for B. & B. (1960), Z. & P. (1969) and
ABBOTT (1974).
Oliva circinata Marrat, 1871 (gv., under ©.
Jfulgurator) was ©. sayana Ravenel, 1834 for ABBOTT
(1974).
Oliva scripta Lamarck, 1811.
Oliva scripta Lamarck, 1811: 315, sp. no. 21; Encycl.
PI. 362, fig .4.
Oliva caribaeensis Dall & Simpson, 1901: 391, PI.
56, fig. 9.
Oliva trujilloi Clench, 1938: 109-113, PI. 9.
Oliva (Cariboliva) scripta venezuelana Petuch &
Sargent, 1986: 71, PL 4, figs. 20, 21.
SIZE: to over 50 mm.
SHAPE: cylindrical.
SPIRE: low conical, telescopic. Spire callus semi-
convex to convex, covering from half to whole
whorl. Colour beige, without ornamentation.
CHANNEL: extremely wide.
SUBCHANNEL PATTERN: fasciculated, variable.
SHELL BACKGROUND: pale beige, occasionally
whitish.
COLOUR PATTERN: network of fine strokes, forming
tents and triangles of variable size. This is overlaid
by fine dark brown markings sometimes arranged
in axial series, concentrated in two spiral bands.
These often contain long fine strokes or chevrons.
Near the shoulder, the angle of these dark chevrons
coincides with that of the fine strokes pattern. In
many specimens, the angle of the fine strokes
pattern becomes progressively larger as one goes
towards the base, while the angle of the dark
chevrons remains nearly constant (see PL. 3, FIG.
3).
COLUMELLA: mostly with coarse plaits over all the
length, often with white enamel.
FASCIOLE: white to faint beige.
SUPRAFASCIOLAR BAND: lower zone with parallel
fine strokes of variable shape; upper zone with
parallel, axial strokes.
30
Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
APERTURE: greyish white, sometimes faint purplish,
rarely violet. Inner lip margin dark in specimens
with sharp lip.
PROTOCONCH: medium large.
Diagnosis. Readily distinguished from all other Oliva
species by its extremely wide channel.
Distribution: From Florida to Brazil.
SYNONYMY.
Oliva scripta Lamarck, 1811. As for all Lamarck's
Oliva species, the type material could not be located.
The original figure is reproduced in PL. 5, FIG. 5. O.
scripta Lamarck, 1811 is the valid name (see
GREIFENEDER, DUCHAMPS & TURSCH 1995) of an
objective species (see TURSCH & HUART 1990), in
agreement with the conclusions of ABBOTT (1974)
and P. & S. (1986). This name has been applied by Z.
& P. (1969) and W. & A. (1978) to another species
with an Indo-Pacific distribution.
The following names are synonyms or designate
local forms:
Oliva caribaeensis Dall & Simpson, 1901. The
holotype (see PL. 5, FIG. 6) (previously 1llustrated by
KAICHER 1989, Part IV, card no. 5526) could in no
way be distinguished from ©. scripta Lamarck, 1811,
of which it is a subjective junior synonym. This is
in agreement with the conclusions of W. & A. (1978),
OLD (cited in Z. & P. 1969), ABBOTT (1974) and
Diaz & PUY ANO (1994). It was a valid species for Z.
& P. (1969), who state that in Puerto Rico, solid color
intergrades link this taxon with ©. reticularis
Lamarck, 1811 -not confirmed on the solid coloured
Portorican specimens examined by us.
Oliva trujilloi Clench, 1938. The holotype (see PL. 5,
FIG. 7) (previously illustrated by KAICHER 1988, Part
IT, card no. 5200) and the six paratypes examined
(see PL. 5, FIG. 8 and FIG. 9) are dead shells that
could not be distinguished from ©. scripta Lamarck,
1811 except for their reddish colouration, commonly
seen in old shells that have been long exposed to
sunlight. ©. trujilloi Clench, 1938 is a subjective
junior synonym of ©. scripta Lamarck, 1811, in
agreement with the conclusions of B. & B. (1960, as
O. caribaeensis), W. & A. (1978), OLp cited in Z. &
P. (1969), ABBOTT (1974), P. & S. (1986) and DIAZ
& PUY ANO (1994).
Oliva scripta venezuelana Petuch & Sargent, 1986.
The holotype (see PL. 5, FIG. 10) (previously
illustrated by KAICHER 1988, Part IIL, card no. 5199)
is linked by an unbroken series of intergrading
specimens (Shells with very similar, nebulose colour
pattern are found in Honduras) to other populations
of O. scripta Lamarck, 1811. It is, at best, a local
form of ©. scripta Lamarck, 1811, in agreement with
the conclusions of DIAZ & PUY ANO (1994).
The following names have been mistakenly or
questionably considered as synonyms:
Oliva leucostoma d'Orbigny, 1842 (belonging to the
genus Olivancillaria according to B. & B., 1960) was
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
considered by W. & A. (1978) to be a probable
synonym of ©. scripta Lamarck, 1811. Even if O.
leucostoma d'Orbigny, 1842 belonged to the genus
Oliva, it would be a younger homonym of ©.
leucostoma Duclos, 1840.
Oliva jamaicensis Marrat, 1867, [a local variety of O.
Jfulgurator (Rôüding, 1798)] was considered by Z. &
P. (1969) to be a possible synonym, and by ABBOTT
(1974) to be a synonym of ©. caribaeensis Dall &
Simpson, 1901.
Oliva graphica Marrat, 1870 [a local variety of O.
Julgurator (Rôding, 1798)] was considered by
ABBOTT (1974) to be a possible synonym of ©.
scripta Lamarck, 1811.
Oliva porcea Marrat, 1870 [a local variety of O.
Jfulgurator (Rôding, 1798)] was considered by
ABBOTT (1974) to be a possible synonym of ©.
scripta Lamarck, 1811.
Oliva spicata (Rôüding, 1798).
Porphyria spicata Rôding, 1798: 35, sp. no. 423.
Porphyria arachnoidea (Rôding, 1798): 36, sp. no.
450.
Oliva araneosa Lamarck, 1811: 315, sp. no. 19.
Oliva venulata Lamarck, 1811: 313, sp. no. 13;
Encycl. PI. 361, fig. 5.
Oliva oniska Duclos, 1845: 19, PL. 32, figs. 7-9.
Oliva pindarina Duclos, 1840: PI. 16, figs. 7-8;
1845: 19.
Oliva timoria Duclos, 1840: PI. 17, figs. 11-13;
Illustr. Conch.: 19 (pars).
Oliva subangulata Philippi, 1848: PI. 1, fig. 2.
Oliva cumingii Reeve, 1850: PI. 11, figs. 19, à, b.
Oliva ligneola Reeve, 1850: sp. no. 57, PI. 21, fig.
57.
Oliva melchersi Menke, 1851: 24.
Oliva intertincta Carpenter, 1857: 465.
Oliva violacea Marrat, 1867: 213.
Oliva brunnea Marrat, 1870: 7, sp. no. 24, figs. 54,
59.
Oliva punctata Marrat, 1870: sp. n° 6, P1.2, figs.
12-13.
Oliva fuscata Marrat, 1870: sp. n° 8, figs. 20-22.
Oliva spicata var. hemphilli Ford in Johnson, 1911:
122
Oliva spicata var. perfecta Johnson, 1911: 122.
Oliva rejecta Burch & Burch, 1962: 165.
Oliva ionopsis Berry, 1969: 163-64.
Oliva (Strephona) radix Petuch & Sargent, 1986:
143, P1.26, figs.20-23.
Oliva subangulata corteziana Petuch & Sargent,
1986: 150, PI. 29, figs. 7-10.
Description.
Local populations are very homogeneous. The ranges
of variations here under describe differences between
populations.
SIZE: from about 25 mm to over 65 mm.
Oliva spicata (Rôding,
SHAPE: very variable, from elongated fusiform to
inflated fusiform. Cylindrical forms do not seem to
occur. Strong tendency to inflated, angular
shoulders (biconical appearance).
SPIRE: conical, rather elevated. Spire callus rarely
covering more than half of whorl, colour from
white, beige, purplish to brown, often darker at
upper margin. Callus often presents oblique dark
strokes.
CHANNEL: of medium width.
SUBCHANNEL PATTERN: fasciculated, very variable.
Not seen in albinistic, melanistic and in unicoloured
specimens.
SHELL BACKGROUND: from white to cream, to yellow,
to gray.
COLOUR PATTERN: tents and chevrons of very
variable size, width and intensity. Colour of
markings varies from reddish to brown or black.
Start of chevrons often marked with small blotch.
Two faint large spiral bands are formed by local
reinforcement of the pattern. Some populations are
nearly devoid of colour pattern and have overall
golden or whitish appearance. Overlay of additional
uniform brown colour often covers the whole shell,
but the Caribbean "hifasciata pattern" is absent (or
very rare).
COLUMELLA: very variable. Colour from white to
greyish, io pale purple.
FASCIOLE: mostly uniform white to greyish, purplish
or brownish. Lower part often suffused with colour.
Ridges often marked with light brown to dark
purplish brown.
SUPRAFASCIOLAR BAND: very variable.
APERTURE: from white to beige, to greyish blue and
purple. Inner margin of lip: variable, from aperture
colour to continuous or interrupted darker
markings.
PROTOCONCH: large to very large.
Diagnosis. The large values of the protoconch
character RESS and the small NW (number of
nuclear volutions) (see Fig. 8, 11-18) set ©. spicata
(Rôding, 1798) apart from all other Oliva species,
excepted ©. fulgurator (Rôding, 1798). In most
cases, it differs from ©. fulgurator (Rôding, 1798) by
much weaker spiral bands, (the Caribbean "bhifasciata
pattern" is absent or very rare), by the presence of
oblique strokes on the spire callus and by the
presence of coloured fasciolar ridges.
Distribution. Widely distributed in the Panamic
region, from the Gulf of California to Ecuador.
SYNONYMY.
1798), based upon the
acceptable figures 509 and 510 of MARTINI (see PL.
10, FIG. 5), is the valid name (see TURSCH,
DUCHAMPS & GREIFENEDER 1994) for an objective
species (see $ 4.1) . This is agreement with the
conclusions of Z. & P. (1969), W. & A .(1978), P. &
S. (1986) and. For a colour variation see JACKSON
(1991).
31
APEX 13(1-2): 1-61, 20 avr. 1998
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
The following names are synonyms or designate
local forms:
"Oliva arachmoidea” (Rôding, 1798) in W. & A.
(1978) is a printing error for ©. arachnoidea
(Rôding, 1798) (q.v.).
Oliva arachnoidea (Rôding, 1798) is an objective
junior synonym of ©. spicata (Rôding, 1798) (see
TURSCH, DUCHAMPS & GREIFENEDER 1994), in
agreement with B. & B. (1960), Z. & .P (1969), KEEN
(1971), ABBOTT (1974), W. & A. (1978) and P. &S.
(1986).
Oliva araneosa Lamarck, 1811. This is an objective
junior synonym of ©. spicata (Rôding, 1798) (see
TURSCH, DUCHAMPS & GREIFENEDER 1994), in
agreement with B. & B. (1960), Z. & P. (1969), KEEN
(1971), ABBOTT (1974), W. & A. (1978) and P. &S.
(1986). It was a colour form of the same for TERZER
(1996).
"Oliva aranersa Lamarck, 1811” (auct) in W. & A.
(1978) is a printing error for ©. araneosa Lamarck,
1811.
Oliva venulata Lamarck, 1811. This is an objective
junior synonym of ©. lifterata (Rôding, 1798) (see
GREIFENEDER, DUCHAMPS & TURSCH 1995), which
is itself a nomen dubium (see TURSCH, DUCHAMPS &
GREIFENEDER 1994). This was a valid species for
DucLos (1845: 25; PI. 17, figs. 5, 6; PI. 22, figs. 19,
20; PI. 33, fig.11, illustrating rather dissimilar shells)
and for P. & S. (1986). It was a form of ©. spicata
(Rôding, 1798) for B. & B. (1960), Z. & P. (1969),
KEEN (1971), ABBOTT (1974) and W. & A. (1978). It
was a separate species for TERZER (1996).
Oliva timoria Duclos, 1840. The type material is
heterogeneous. One of the syntypes (see PL. 10, FIG.
1) 1s very similar to ©. radix Petuch & Sargent, 1986,
so ©. timoria Duclos, 1840 is (pars) a subjective
junior synonym of ©. spicata (Rôüding, 1798).
“Oliva onisca Duclos, 1844” (auct) is a misspelling
for ©. oniska Duclos, 1845. This was ©. fulgurator
(Rôding, 1798) for B. & B. (1960).
Oliva oniska Duclos, 1845. The type series consists in
6 mottled specimens and 3 brown specimens,
including the figured syntype (see PL. 11, FIG. 4)
(illustrated by KAICHER 1989, Part IV, card no.
5473). The type material could not be separated from
the “spicata complex” morphological continuum,
being linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, this is here considered as a
subjective junior synonym of ©. spicata (Rôding,
1798), in agreement with Z. & P. (1969). This was O.
fulgurator (Rôding, 1798) for W. & A. (1978); a
subspecies of ©. tisiphona Duclos, 1845 (a nomen
dubium) for P. & S. (1986).
Oliva pindarina Duclos, 1840. The type material,
including the figured syntype (see PL. 10, FIG. 2)
(illustrated by KAICHER 1989, Part IV, card no.
5523), could not be separated from the “spicata
complex” morphological continuum, being linked to
32
all other conspecific forms by many intergrading
populations. In the absence of valid arguments to the
contrary, this is here considered as a local form of ©.
spicata (Rôding, 1798), in agreement with B. & B.
(1960), Z. & P. (1969), KEEN (1971), ABBOTT (1974)
and W. & A. (1978). This was a colour form of ©.
venulata Lamarck, 1811 for P. & S. (1986) and for
TERZER (1996). Similar to some forms of ©.
fulgurator
Oliva subangulata Philippi, 1848. This is represented
in BM(NH) by two specimens, with the label "type"
fallen off. One of the specimens bears traces of glue
and is most probably the holotype. This shell (see PL.
11, FIG. 6), with a slight shoulder bulge, could not be
separated from the “spicata complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. In the
absence of valid arguments to the contrary, this is
here considered as a local form of ©. spicata
(Rôding, 1798), in agreement with Z. & P. (1969),
KEEN (1971), ABBOTT (1974) and W. & A. (1978).
This was a valid species for P. & S. (1986) and for
TERZER (1996).
Oliva cumingii Reeve, 1850. The three syntypes (see
PL. 11, FIG. 1) (one is illustrated by KAICHER 1989,
Part IV, card no. 5551) with the label "Gulf of
California. H. Cuming colin." have a banded colour
pattern and are somewhat bulging. The type material
could not be separated from the “spicata complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of ©.
spicata (Rôding, 1798), in agreement with Z. & P.
(1969), KEEN (1971), ABBOTT (1974) and W. & A.
(1978). It was a subspecies of ©. venulata Lamarck,
1811 for P. & S. (1986), a form of the same for
TERZER (1996).
Oliva ligneola Reeve, 1850. One of the two
heterogeneous syntypes (see PL. 10, FIG. 3) is a
specimen of ©. tigrina fallax Johnson, 1910, for
which ligneola Reeve, 1850 would thus be an earlier
name. The other one is a bleached shell that could not
be separated from the “spicata complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, ©.
ligneola Reeve, 1850 (pars) is here considered as yet
another subjective junior synonym of ©. spicata
(Rôding, 1798). This was a nomen dubium for W. &
A. (1978).
Oliva melchersi Menke, 1851. There is no type
material and no original illustration. The original
description says that the shell is angular, ash and
flesh in colour, with yellowish ("ochroleucus"),
triangular blotches and brown dots. The shoulder
presents a pattern of lines in bundles("American
pattern"). Lower whorls of spire uniformly flesh
coloured, with brown spots above; lip sharp, with
black maculations. The above description, together
with the type locality (Mazatlan, Mexico) and the
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
mention by the author that it could be a freak of O.
venulata Lamarck, 1811, indicates this is a
subjective junior synonym of O. spicata (Rôding,
1798), in agreement with B. & B. (1960), Z. & P.
(1969), KEEN (1971), ABBOTT (1974), and W. & A.
(1978). It was a subspecies of ©. spicata (Rôding,
1798) for P. & S. (1986) and for TERZER (1996).
Oliva intertincta Carpenter, 1857. The two
homogeneous syntypes USNM 716187 (see PL. 11,
FIG. 9) are dead, discoloured, juvenile specimens of
O. spicata (Rôding, 1798). So are the three additional
syntypes at BM(NH) (see KEEN 1968). AII have a
brown fasciole and the second whorl of the
protoconch is more inflated than usual, both
characters intergrading with ©. spicata (Rüding,
1798), of which this is here considered to be a
subjective junior synonym. This is in agreement
with the conclusions of B & B (1960), Z & P (1969),
KEEN (1971) ABBOTT (1974) W & A (1978), P & S
(1986) and TERZER (1996).
Oliva violacea Marrat, 1867. The holotype (see PL. 11,
FIG. 8) (illustrated by KAICHER 1989, Part IV, card
no. 5479) could not be separated from the “spicata
complex” morphological continuum, being linked to
all other conspecific forms by many intergrading
populations. In the absence of valid arguments to the
contrary, this is here considered as yet another local
variety of O. spicata (Rôding, 1798), in agreement
with B. & B. (1960), KEEN (1971), ABBOTT (1974),
Z. & P. (1969) and W. & A. (1978). This was a valid
species for TERZER (1996) and for P. & S. (1986), but
O. violacea ; Petuch & Sargent (not Marrat, 1867),
1986, depicted in the Atlas, PI. 30, figs. 20, 21 differs
very much from the type specimen.
Oliva brunnea Marrat, 1870. The brown holotype (see
PL. 11, FIG. 3) (illustrated by KAICHER 1989, Part IV,
card no. 5573) is nearly identical to one of the
syntypes of Oliva fuscata Marrat, 1870 (q.v., see PL.
11, FIG. 2). It could not be separated from the
“spicata complex” morphological continuum, being
linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, this is here considered as
yet another local variety (and colour form) of ©.
spicata (Rôding, 1798), in agreement with B. & B.
(1960) and W. & A. (1978). It was a subspecies of
the Atlantic taxon ©. jamaicensis Marrat, 1870 for P.
& S. (1986). This taxon is indeed one of the cases of
striking resemblance between populations of ©.
Julgurator (Rôding, 1798) and populations of ©.
spicata (Rôding, 1798) (see $ 5.4). Some specimens
from Paranagua, Venezuela are extremely similar to
their Pacific counterparts.
Oliva punctata Marrat, 1870. This name is available
because ©. punctata (Rôding, 1798) is a nomen
nudum (see TURSCH, DUCHAMPS & GREIFENEDER
1994). According to TOMLIN (in FORD, 1953) the 8
specimens at MCM are not types because the largest
specimen (said to be 33 mm x 16 mm) is not large
enough. But according to MCMILLAN (1985), the two
syntypes are the originals of Marrat's figures. In any
case, these specimens (see PL. 10, FIG. 4) could not
be separated from the ‘“spicata complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
In the absence of valid arguments to the contrary, this
is here considered as yet another local variety of O.
spicata (Rôding, 1798), in agreement with Z. & P.
(1969), KEEN (1971) and ABBOTT (1974). This was
O. spicata var. venulata Lamarck, 1811 for B. & B.
(1960), O. rejecta Burch & Burch, 1962 for P. & S.
(1986) and for TERZER (1996). "punctulata Johnson,
1911" is a misquotation in B & B (1960) and W & A
(1978) for punctata Marrat, cited in Johnson (1911:
122) under spicata.
Oliva fuscata Marrat, 1870. The type material consists
of three syntypes [Note: TOMLIN (in FORD, 1953)
mentions six syntypes]. Two are mottled (one is
figured PL. 11, FIG. 7); the last (see PL. 11, FIG. 2) is
brown and nearly identical with the type of O.
brunnea Marrat, 1870. The type material could not be
separated from the “spicata complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. In the
absence of valid arguments to the contrary, this is
here considered as yet another local variety of ©.
spicata (Rôding, 1798), in agreement with B. & B.
(1960), Z. & P. (1969), KEEN (1971), ABBOTT
(1974), W. & A. (1978), P. & S. (1986) and TERZER
(1996).
Oliva spicata var. hemphilli Ford in Johnson, 1911.
The five syntypes (see PL. 10, FIG. 8) could not be
separated from the “spicata complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. In the
absence of valid arguments to the contrary, this is
here considered as yet another local variety of ©.
spicata (Rôding, 1798), in agreement with Z. & P.
(1969), KEEN (1971), ABBOTT (1974) and W. & A.
(1978). This was a colour form of ©. venulata
Lamarck, 1811 for P. & S. (1986) and for TERZER
(1996).
Oliva spicata Var. perfecta Johnson, 1911. The three
homogeneous, dark syntypes (the largest is figured
here PL. 11, FIG. 5) could not be separated from the
“spicata complex” morphological continuum, being
linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, this 1s here considered as
yet another colour form of ©. spicata (Rôding,
1798), in agreement with B. & B. (1960), Z. & P.
(1969) and KEEN (1971). This was ©. fuscata Marrat,
1870 for W. & A. (1978), O. venulata Lamarck, 1811
for P & S (1986) and TERZER (1996).
Oliva rejecta Burch & Burch, 1962 is a specially
interesting case, as this is the only report known to us
of syntopic coexistence of two forms within the
"spicata complex" (an information contradicted by
recent field observations, see $ 4.3.2). The examined
paratypes at USNM (see PL. 10, FIG. 10) (illustrated
by KAICHER 1989, Part IV, card no. 5527)and
MNAN (see PL. 10, FIG. 9) could not be separated
33
APEX 13(1-2): 1-61, 20 avr. 1998
from the “spicata complex” morphological
continuum, being linked to all other conspecific
forms by many intergrading populations. The
reported differences -base of columella light purple
instead of white, D/H 0.42 (calculated from reported
measurements of two specimens only; 0.43 measured
on paratypes) instead of 0.49- fall well within the
variation range of the “spicata complex”. The other
reported difference -brittleness of the shell, making it
unsuitable for producing Indian artifacts- and
reported differences in the X-Ray diffraction pattern
of the shell (DONOHUE & HARDCASTLE 1962) could
not be tested by us, as this would entail destruction of
type material. These could certainly represent
interpopulation variation. In the absence of valid
arguments to the contrary, this is here considered as
yet another local variety of ©. spicata (Rôding,
1798), in agreement with the conclusions of Z & P
(1969) KEEN (1971) ABBOTT (1974). It was a valid
species (Oliva punctata Marrat, 1870) for P. & S.
(1986) and for TERZER (1996). This form comes from
La Paz, Baja California, Mexico, on tide flats.
Oliva ionopsis Berry, 1969: 163-64. The holotype (see
PL. 11, FIG. 10) (previously illustrated in HERTZ
(1984: 37, Fig. 89) and the seven paratypes could not
be separated from the “spicata complex”
morphological continuum, being linked to all other
conspecific forms by many intergrading populations.
For instance, the blue aperture (to which the name
refers) is also found in the varieties violacea Marrat,
1867, rejecta Burch and Burch, 1962 as well as in
many unnamed forms. In the absence of valid
arguments to the contrary, this is here considered as
yet another local variety of ©. spicata (Rôding,
1798), in agreement with KEEN (1971) and TERZER
(1996). It was considered a valid species by ABBOTT
(1974), W. & A. (1978) and P. & S. (1986). The type
locality is Bahia de las Palmas, Baja California, in
10-33 fms.
Oliva radix Petuch & Sargent, 1986. The holotype (see
PL. 10, FIG. 6) (previously 1illustrated by KAICHER
1989, Part IV, card no. 5486) has an angulose shape
that somewhat reminds of some bulgy forms of ©.
polpasta Duclos (a species broadly co-occurring in
Ecuador, see KEEN, 1971) but lacks the characteristic
subchannel "cogwheel" pattern of this species (see
under ©. polpasta). Shells of ©. spicata (Rôding,
1798) with rather similar shapes (but less bulgy)
occur in Western Mexico. The protoconch
measurements (RES5= 1.58, NW=2.50, see Fig. 3)
and (PATI8/NW = 0.4, Ipro = 0.82, see Fig. 6) also
indicate it is a local variety of O. spicata (Rôding,
1798).
Oliva subangulata corteziana Petuch & Sargent, 1986.
In spite of its more bulging body whorl (see PL. 10,
FIG. 7), the holotype could not be separated from the
“spicata complex” morphological continuum, being
linked to all other conspecific forms by many
intergrading populations. In the absence of valid
arguments to the contrary, this is here considered as
yet another local variety of O. spicata (Rüding,
34
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
1798). This form comes from the Northern end of the
Gulf of California. This was a valid subspecies of ©.
subangulata Philippi, 1848 for TERZER (1996).
The following names have been mistakenly (or very
questionably) considered as synonyms:
Oliva harpularia Lamarck, 1811 is a nomen dubium
(see GREIFENEDER, DUCHAMPS & TURSCH 1995).
This was ©. spicata (Rôding, 1798) for B. & B.
(1960) and W. & A. (1978), a colour form of ©.
venulata Lamarck,1811 for P. & S. (1986), a possible
synonym of ©. pindarina Duclos, "1835" for TERZER
(1996).
Oliva ustulata Lamarck, 1811 is another nomen
dubium (see GREIFENEDER, DUCHAMPS & TURSCH
1995). This was a synonym of ©. reticularis
Lamarck, 1811 for DUCLOS (1845, p. 16), ©. spicata
(Rôding, 1798) for B. & B. (1960); a form of ©.
spicata (Rôding, 1798) for Z. & P. (1969), possibly
O. fuscata Marrat, 1870 for W. & A. (1978); a colour
form of ©. venulata Lamarck,1811 for P. & S. (1986)
and for TERZER (1996).
Oliva oriola ; Duclos (not Lamarck, 1811), 1835: PI.
10, figs. 1, 2, text in ///ustr. Conch.: 15. This was O.
spicata (Rôding, 1798) for B. & B. (1960), Z. & P.
(1969), KEEN (1971), ABBOTT (1974) and W. & A.
(1978). This name has already been treated under ©.
fulgurator (Rôding, 1798) (g.v.).
Oliva obesina Duclos, 1840. This was ©. spicata
Rôding for B. & B. (1960), Z. & P. (1969), KEEN
(1971), ABBOTT (1974) and W. & A. (1978); à
separate Panamic species for TERZER (1996). This
name has already been treated under ©. fulgurator
(Rôding, 1798) (q.v.).
Oliva ispida (Link) (not Rôding, 1798), 1807. This was
O. spicata (Rôding, 1798) for B. & B. (1960). This
name has already been treated under ©. fulgurator
(Rôding, 1798) (q.v.).
Oliva oblonga Marrat, 1867. This was ©. spicata
(Rôding, 1798) for B. & B. (1960), KEEN (1971) and
W. & A. (1978). This name has already been treated
under ©. fulgurator (Rüding, 1798) (q.v.).
Oliva graphica Marrat, 1870. This was ©. spicata
(Rôding, 1798) for B. & B. (1960). This name has
already been treated under ©. fulgurator (Rôding,
1798) (q.v.).
“Oliva oblongata Marrat, 1870” (auct). This was O.
spicata (Rôding, 1798) for Z. & P. (1969), W. & A.
(1978), ABBOTT (1974) and P. & S. (1986). This
name has already been treated under ©. fulgurator
(Rôding, 1798) (q.v.).
Oliva porcea Marrat, 1870. This was ©. spicata
(Rôding, 1798) for B. & B. (1960). This name has
already been treated under ©. fulgurator (Rôding,
1798) (q.v.).
One name is still unsolved:
Oliva schumacheriana Beck in Gray (1858: 46)
(original name: Strephona schumacheriana), cited by
B. & B. (1960) as "Beck 1858 Cal. Proc. 1858, p.46")
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
could not be tracked. The only indication in Gray is
"Front of pillar lip brown. Hab. California". This
possibly refers to the ridges of the fasciole, often
marked with brown in ©. spicata (Rôding, 1798).
Oliva spicata deynzerae Petuch & Sargent,
1986.
Oliva (Strephona) spicata deynzerae Petuch & Sargent,
1986: 149, PI. 28, figs. 12, 13.
Description.
SIZE: to over 55 mm.
SHAPE: elongated fusiform.
SPIRE: conical, slightly telescopic. Spire callus gray
brown to purple, covering up to 2/3 of whorl, with
dark brown oblique strokes.
CHANNEL: medium.
SUBCHANNEL PATTERN: fasciculated, of variable
intensity.
SHELL BACKGROUND: whitish.
COLOUR PATTERN: Network of fine tent marks
(giving bluish-gray aspect) is overlaid with dark
brown little blotches, especially concentrated to
form two large spiral bands, in which are found
strokes, chevrons and ziczacs.
COLUMELLA: with sharp plaits over all the length.
Colour purplish white.
FASCIOLE: same colour as columella, with smoky
base and upper margin purple.
SUPRAFASCIOLAR BAND: two similar zones of
puncticulated brown marks.
APERTURE: beige-cream.
PROTOCONCH: medium large.
Diagnosis. Spire more telescopic than ©. spicata.
Double spiral band more conspicuous.
Distribution. Known only from the Cocos Islands,
Costa Rica.
Oliva spicata deynzerae Petuch & Sargent, 1986. The
holotype (see PL. 4, FIG. 5) and the six topotypes that
were examined could be separated from all the
remainder of the "©. spicata complex" (see $ 5. 3 and
Fig. 18). This is therefore considered as a subspecies
of Oliva spicata (Rôding, 1798), in agreement with P.
& S. (1986) and TERZER (1996). All specimens
examined had a very similar colour pattern, curiously
reminiscent of that of the sympatric species Oliva foxi
Stingley, 1984 (local crypsis pattern?). The
subspecies status rests on a rather small sample (see $
5. 3) and would have to be modified if another form
of O. spicata were to be discovered in the Cocos Is.
Acknowledgements.
We are most grateful to Dr. G. ROSENBERG and Dr.
E. S. GILMORE (Academy of Natural Sciences,
Philadelphia), Dr. W. EMERSON (American Museum of
Natural History, New York), Dr. W. PONDER and Dr. I.
LOCH (Australian Museum, Sydney), Dr. R. JANSSEN
(Forschungsinstitut und Naturmuseum Senckenberg,
Frankfurt), Dr. I. WALLACE, (Merseyside County
Museum, Liverpool), Dr. Y. FINET (Muséum d'Histoire
Naturelle, Genève), Dr. Ph. BOUCHET and Mme. A.
TILLIER (Muséum National d’ Histoire Naturelle,
Paris), Prof. K J. BOSS (Museum of Comparative
Zoology, Harvard University), Dr. M.G. HARASEWYCH
and Ms. R. N. GERMON [National Museum of Natural
History (Smithsonian Institution), Washington], Dr. H.
CHANEY (Santa Barbara Museum of Natural History)
and Ms. K. WAY (The Natural History Museum,
London), for allowing us to study type material and
specimens from historical collections.
Mr. André BOssSUYT (Wervik, Belgium), Mrs.
Mercedes CAMACHO (La Paz, Mexico), Mr. Venerado
CANTERO (Cardenas, Cuba), Mr. Alex CORBO
(Brussels, Belgium), Dr. Juan DIAZ (Instituto de
Investigaciones Marinas de Punta de Betin, Santa
Marta, Colombia), Mr. Ralph DUCHAMPS (Brussels,
Belgium), Dr. José ESPINOSA (Instituto de
Oceanologia, Academia de Ciencias de Cuba), Prof.
Esteban Fernando FÉLIX PICO (CICIMAR, La Paz,
Mexico), Mr. Raül FERNANDEZ-GARCÉS (Cienfuegos,
Cuba), Mr. Luc GERMAIN (now in Bujumbura,
Burundi), Mr. Jose Luis Peña GONZALEZ (Cardenas,
Cuba), Dr. Yuri KANTOR (Severtzov Institute,
Moscow, Russia), Prof. Alan KOHN (University of
Washington, Seattle, Washington, USA), Dr. Harry G.
LEE (Jacksonville, Florida, USA), Mr. Mauricio
Andrade LIMA (Recife, Brazil), Mrs. Mabel
MARTINEZ CRUZ (Santa Marta, Veradero, Cuba), the
late Dr. E. MATTHEWS (Titusville, Florida, USA), Mr.
Brian PARKINSON (Auckland, New Zealand), Prof.
Pablo E. PENCHASZADEH (Universidad Simon Bolivar,
Caracas, Venezuela), Mr. Jean PIERRET (now in Palo
Alto, California, USA), Mr. Guido POPPE (Brussels),
Mr. Jesus Antonio RAMIREZ HERNANDEZ (Cartagena,
Colombia), Mr. Colin REDFERN (Boca Raton, Florida,
USA), Mr. André VASSART (s/s La Boudeuse), Dr.
Pablo RAMIREZ VILLAROEL (Universidade de Oriente,
Porlamar, Venezuela), Prof. Eli RIOS (Museu
Oceanogräfico, Rio Grande, RS, Brazil), Dr. Jacques
SENDERS (Brussels, Belgium), Mr. Christian VAN
OSSELAER (Brussels, Belgium), Mr. Olivier VRONEN
(Brussels, Belgium) and Mr. Bob WoRK (South Miami,
Florida, USA) have been of great assistance by either
loaning specimens, providing valuable advise and
information, helping with computers and programming
problems, taking part in field work, or correcting the
manuscript. We are most grateful to all.
We are especially indebted to Mrs. Kathie WAY
(BMNH, London) and Mr. Antoine LIEVROUW
(IRSNB, Brussels) for their kind cooperation in
tracking old literature. We thank two unknown referees
for their remarks and corrections.
We are grateful to the Fonds National de la
Recherche Scientifique (F.N.R.S.) and BIOTEC, S.A.
35
APEX 13(1-2) 1-61, 20 avr. 1998
8. INDEX TO NAMES (objective species in bold)
“alba Lamarck": misquotation in Gray (1858).
“aldina” Duclos: misspelling for ©. aldinia Duclos,
1845.
aldinia Duclos, 1845: subjective junior synonym of ©.
fulgurator (Rôding, 1798).
angulata Lamarck, 1811: objective junior synonym of
O. incrassata (Rôding, 1798).
antillensis Petuch & Sargent, 1986: local variety of ©.
fulgurator (Rôding, 1798).
"arachmoidea” (Rèding, 1798) (auct.): printing error
for ©. arachnoidea (Rôding, 1798).
arachnoidea (Rôding, 1798): objective junior synonym
of O. spicata (Rôding, 1798).
araneosa Lamarck, 1811: objective junior synonym of
O. spicata (Rôding, 1798).
aranersa" Lamarck, 1811 (auct.): printing error for
©. araneosa Lamarck, 1811.
bahamasensis Petuch & Sargent, 1986: local variety of
O. fulgurator (Rôding, 1798).
barbadensis Petuch & Sargent, 1986: local variety of
O. fulgurator (Rôding, 1798).
bewleyi Marrat, 1870: local variety of ©. fulgurator
(Rôding, 1798).
bifasciata Küster in Weinkauff, 1878: colour form of
O. fulgurator (Rôding, 1798).
bifasciata jenseni Petuch & Sargent, 1986: local
variety of ©. fulgurator (Rôding, 1798).
bifasciata sunderlandi Petuch, 1987: local variety of ©.
fulgurator (Rôding, 1798).
bollingi Clench, 1934: see reticularis bollingi Clench,
1934.
broderipi Ducros de St. Germain, 1857: local variety of
O. fulgurator (Rôding, 1798).
brunnea Marrat, 1870: local variety of O. spicata
(Rôüding, 1798).
bullata Marrat, 1871: local variety of ©. fulgurator
(Rôding, 1798).
caribaeensis Dall & Simpson, 1901: subjective junior
synonym of ©. scripta Lamarck, 1811.
circinata Martyn, 1789:nomen nudum (in a rejected
work).
circinata Marrat, 1871: local variety of ©. fulgurator
(Rôding, 1798).
circinata var. citrina Johnson, 1911: colour form of ©.
sayana Ravenel, 1834.
circinata tostesi Petuch, 1987: local variety of O.
Jfulgurator (Rôding, 1798).
citrina Johnson, 1911: see circinata var. citrina
Johnson, 1911
contoyensis Petuch, 1988: local variety of ©. fulgurator
(Rôding, 1798).
corteziana Petuch & Sargent, 1986: see subangulata
corteziana Petuch & Sargent, 1986.
cribraria Marrat, 1833? : local variety of O. fulgurator
(Rôding, 1798).
36
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
cumingii Reeve, 1850: local variety of ©. spicata
(Rôding, 1798).
deynzerae Petuch & Sargent, 1986: see spicata
deynzerae.
“diaphana Duclos 1835” (auct.): misquotation.
drangai Schwengel, 1951: local variety of ©.
Julgurator (Rôüding, 1798).
ernesti Petuch 1990: local variety of ©. fulgurator
(Rôding, 1798).
figura Marrat, 1870: local variety of ©. fulgurator
(Rôding, 1798).
finlayi Petuch & Sargent, 1986: local variety of ©.
fulgurator (Rôding, 1798).
Jormosa Marrat, 1870: local variety of ©. fulgurator
(Rôding, 1798).
foxi Stingley, 1984: valid.
fulgurator (Rôding, 1798): valid.
Juscata Marrat, 1870: local variety of ©. spicata
(Rôüding, 1798).
Jusiformis Lamarck, 1811: objective junior synonym of
©. fulgurator (Rôding, 1798).
goajira Petuch & Sargent, 1986: local variety of ©.
fulgurator (Rôding, 1798).
graphica Marrat, 1870: local variety of ©. fulgurator
(Rôding, 1798).
greenwayae Clench, 1937: see reticularis greenwayae
Clench, 1937.
harpularia Lamarck, 1811: nomen dubium.
hemphilli Ford in Johnson, 1911: see spicata var.
hemphilli Ford in Johnson, 1911.
hepatica ; Marrat (not Lamarck, 1811), 1871: local
variety of ©. fulgurator (Rôding, 1798).
incrassata (Lightfoot in Solander, 1786): valid.
ionopsis Berry, 1969: local variety of ©. spicata
(Rôding, 1798).
intertincta Carpenter, 1857: subjective junior synonym
of O. spicata (Rôüding, 1798)
ispida (Link) (not Rôding, 1798), 1807: objective
junior synonym of ©. fulgurator (Rôding, 1798).
jamaicensis Marrat, 1867: local variety of O.
fulgurator (Rôüding, 1798).
jamaicensis zombia Petuch & Sargent, 1986: local
variety of ©. fulgurator (Rôding, 1798).
jenseni Petuch & Sargent, 1986: see bifasciata jenseni
Petuch & Sargent, 1986.
kerstitchi da Motta, 1985: colour form of ©. polpasta
Duclos, 1833.
ligneola Reeve, 1850 (pars): subjective junior
synonym of ©. spicata (Rüding, 1798); (pars): earlier
name for tigrina fallax Johnson, 1910.
litterata (Rôüding, 1798): nomen dubium.
litterata Lamarck, 1811: junior homonym of O.
litterata (Rôding, 1798).
magdae Petuch & Sargent, 1986: local variety of O.
fulgurator (Rôding, 1798).
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
maya Petuch & Sargent, 1986: local variety of O.
Jfulgurator (Rôding, 1798).
melchersi Menke, 1851: subjective junior synonym of
O. spicata (Rôding, 1798).
memnonia Duclos, 1845: nomen dubium.
mercatoria Marrat, 1871: local variety of ©. fulgurator
(Rüding, 1798).
nivosa Marrat, 1871: local variety of ©. fulgurator
(Rôding, 1798).
obesina Duclos, 1840: local variety of ©. fulgurator
(Rôding, 1798).
oblonga Marrat, 1867: local variety of O. fulgurator
(Rôding, 1798).
“oblongata Marrat, 1870” (auct): misspelling (and a
date error) for ©. oblonga Marrat, 1867.
olivacea Marrat, 1870: local variety of ©. fulgurator
(Rôding, 1798).
olorinella Duclos, 1835 (pars): local variety of ©.
fulgurator (Rôding, 1798).
olorinella Duclos, 1835 (pars): subjective junior
synonym of ©. oliva (L., 1758).
olssoni Petuch & Sargent, 1986: local variety of ©.
polpasta Duclos, 1833.
“onisca Duclos, 1844” (auct): misspelling for ©. oniska
Duclos, 1845.
oniska Duclos, 1845: subjective junior synonym of ©.
spicata (Rôding, 1798).
oriola ; Duclos (not Lamarck, 1811), 1835: nomen
dubium.
pallida Marrat, 1867: local variety of ©. fulgurator
(Rôding, 1798).
pattersoni, Clench, 1945 4: local form and colour
variety of ©. fulgurator (Rüding, 1798).
perfecta Johnson, 1911: see spicata var. perfecta
Johnson, 1911.
pindarina Duclos, 1840: local variety of O. spicata
(Rôding, 1798).
polpasta Duclos, 1833: valid.
porcea Marrat, 1870: local variety of ©. fulgurator
(Rôding, 1798).
punctata (Rôding, 1798): nomen nudum.
punctata Marrat, 1870: local variety of O. spicata
(Rôding, 1798).
"punctulata Johnson, 1911" (auct.): misquotation.
quersolina Duclos, 1835: subjective junior synonym of
©. atalina Duclos, 1835.
radix Petuch & Sargent, 1986: local variety of ©.
spicata (Rôding, 1798).
reclusa Marrat, 1871: local variety of ©. fulgurator
(Rüding, 1798).
rejecta Burch & Burch, 1962: local variety of ©.
spicata (Rôding, 1798).
reticularis Lamarck, 1811: subjective junior synonym
of ©. fulgurator (Rôding, 1798).
reticularis bollingi Clench, 1934: local variety of O.
fulgurator (Rôding, 1798).
reticularis greenwayae Clench, 1937: local form and
colour variety of ©. fulgurator (Rüding, 1798).
sarasotensis Petuch & Sargent, 1986: see sayana
sarasotensis Petuch & Sargent, 1986.
sargenti Petuch, 1987: local variety of ©. fulgurator
(Rôding, 1798).
sayana Ravenel, 1834: valid.
sayana sarasotensis Petuch & Sargent, 1986: local
form of ©. sayana Ravenel, 1834.
sayana texana Petuch & Sargent, 1986: local form of
O. sayana Ravenel, 1834.
schumacheriana Beck in Gray (1858): unknown to us,
possibly ©. spicata (Rôding, 1798).
scripta Lamarck, 1811: valid.
scripta venezuelana Petuch & Sargent, 1986: local
form of ©. scripta Lamarck, 1811.
sowerbyi Marrat, 1870: subjective junior synonym of
O. atalina Duclos, 1835.
spicata (Rôding, 1798): valid.
spicata deynzerae Petuch & Sargent, 1986: subspecies
of O. spicata (Rôding, 1798).
spicata Var. hemphilli Ford in Johnson, 1911: local
variety of ©. spicata (Rôüding, 1798).
spicata var. perfecta Johnson, 1911: colour form of ©.
spicata (Rôding, 1798).
subangulata Philippi, 1848: local variety of ©. spicata
(Rôding, 1798).
subangulata corteziana Petuch & Sargent, 1986: local
variety of ©. spicata (Rôding, 1798).
sunderlandi Petuch, 1987: see bifasciata sunderlandi
Petuch, 1987.
texana Petuch & Sargent, 1986: see sayana texana
Petuch & Sargent, 1986.
timoria Duclos, 1840 (pars): subjective junior
synonym of ©. fulgurator (Rôding, 1798).
timoria Duclos, 1840 (pars): subjective junior
synonym of ©. spicata (Rôding, 1798).
tisiphona Duclos, 1845: nomen dubium.
tostesi Petuch, 1987: see circinata tostesi Petuch, 1987.
trujilloi Clench, 1938: subjective junior synonym of ©.
scripta Lamarck, 1811.
truncata Marrat, 1867: subjective junior synonym of ©.
polpasta Duclos, 1833.
ustulata Lamarck, 1811: nomen dubium.
venezuelana Petuch & Sargent, 1986: see scripta
venezuelana Petuch & Sargent, 1986.
venulata Lamarck, 1811: objective junior synonym of
O. litterata (Rôding, 1798).
"vermiculata Gray, 1858" (auct.): misquotation and
nomen nudum.
"vermiculata Lamarck": misquotation in Gray (1858).
violacea Marrat, 1867: local variety of ©. spicata
(Rôding, 1798).
zombia Petuch & Sargent, 1986: see jamaicensis
zombia Petuch & Sargent, 1986.
37
APEX 13(1-2): 1-61, 20 avr. 1998
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
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Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
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39
APEX 13(1-2): 1-61, 20 avr. 1998
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
TURSCH, B. & D. GREIFENEDER. 1996. The "“Oliva
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on Olividae XIV. The taxonomic structure of Oliva
oliva (auct.). Apex 7(1): 3-22.
VAN OSSELAER, C., J. BOUILLON, J.M. OU & B.
TURSCH. 1993. Studies on Olividae XVIII. The
Nurenberg.
ZEIGLER, R.F. 1969. Two infrasubspecific forms in
Oliva. The Nautilus 83(1): 14-19.
ZEIGLER, R.F. & H.C. PORRECA. 1969. Olive shells of
the world. Rochester Polychrome Press, Rochester,
N.Y.
PLATE 1.
Examples of subchannel colour pattern in Atlantic and Eastern Pacific Oliva species (see text $ 4.1).
1. O. foxi Stingley, 1984. Cocos Is., Costa Rica.
. ©. foxi Stingley, 1984. Cocos Is., Costa Rica.
. ©. porphynia (Linnaeus, 1758). W. Mexico.
. ©. porphyria (Linnaeus, 1758). Cebaco |., W. Panama.
. ©. polpasta Duclos, 1833. Kobbe Beach, W. Panama.
. ©. polpasta Duclos, 1833. Montijo Bay, W. Panama.
. ©. incrassata (Lightfoot in Solander, 1786). Baja California, W. Mexico.
. ©. incrassata (Lightfoot in Solander, 1786). Baja California, W. Mexico.
. ©. julieta Duclos, 1840. Gubernadora |., W. Mexico.
10. O. julieta Duclos, 1840. Manzanillo, W. Mexico.
© © HO On B © ND
11. O. splendidula Sowerby, 1825. W. Panama.
12. O. splendidula Sowerby, 1825. W. Panama.
13. O. spicata (Rôding, 1798). Baja California, W. Mexico.
14. O. spicata (Rôding, 1798). Baja California, W. Mexico.
15. O. sayana Ravenel, 1834. Marco Beach, Florida, USA.
16. O. sayana Ravenel, 1834. Morgans Pass, Florida, USA.
17. O. scripta Lamarck, 1811. Honduras.
18. O. scripta Lamarck, 1811. Honduras.
19. ©. fulgurator (Rôding, 1798) f. reticularis Lamarck, 1811. E. Panama.
20. O. fulgurator (Rôding, 1798). Aruba.
21. O. fulgurator (Rôding, 1798) f. circinata Marrat, 1871. lIha Bela, Brazil.
22. O. fulgurator (Rôding, 1798) f. circinata Marrat, 1871. Salvador, Brazil.
23. O. spicata deynzerae Petuch & Sargent, 1986. Cocos Is., Costa Rica.
24. O. spicata deynzerae Petuch & Sargent, 1986. Cocos Is., Costa Rica.
25. O. kaleontina Duclos, 1835. Melon I., W. Panama.
26. O. kaleontina Duclos, 1835. Cebaco |., W. Panama.
27. O. undatella Lamarck, 1811. W. Mexico.
28. O. undatella Lamarck, 1811. San Pedro, Ecuador.
29. O. peruviana Lamarck, 1811. Iquique, Chile.
30. ©. peruviana Lamarck, 1811. Iquique, Chile.
31. ©. flammulata Lamarck, 1811. Luanda, Angola.
32. O. flammulata Lamarck, 1811. La Awera, Mauritania.
33. O. flammulata dolicha Locard, 1896. S&o Vicente, Cabo Verde.
O.
flammulata dolicha Locard, 1896. S#o Vicente, Cabo Verde.
ERRATUM
APEX, Vol. 12(4), 20 décembre 1997
Description d’une nouvelle espèce de Costellariidae des Philippines, par E. Guillot de Suduiraut.
Page 119 : lire Vexillum (Costellaria) sauternesense n. sp. au lieu de F. (C.) sauternesi n. sp.
Page 119 : read Vexillum (Costellaria) sauternesense n. sp. instead of V. (C.) sauternesi n. sp.
COLLECTION
Guido T. POPPE
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85, rue des Coteaux Fleuris - 83200 TOULON
Tél. 04 94 92 96 21 - Fax 04 94 22 97 46 Gauguini
TURSCH, GREIFENEDER & HUART Oliva fulgurator and related species APEX 13(1-2): 1-61, 20 avr. 1998
nd EL DS
1. O. foxi 2. O. foxi 3. O. porphyria 4. O. porphyria 6. O. polpasta
7. O. incrassata 8. O. incrassata 9. ©. julieta 10. ©. julieta 11. O. splendidula 12. O. splendidula
14. O. spicata 15. O. sayana 16. O. sayana 17. O. scripta 18. O. scripta
7144 ° P4AMN
le" ME:
+ we + : 3
19. O. fulgurator 20. ©. fulgurator 21. O. fulgurator 22. ©. fulgurator 23. ©. spicata 24. O. spicata
f. reticularis f. circinata f. circinata deynzerae deynzerae
25. O. kaleontina 26. O. kaleontina 27. O. undatella 28. ©. undatella 29. ©. peruviana 30. O. peruviana
31. O. flammulata 32. O. flammulata 33. O. flammulata 34. O. flammulata
dolicha dolicha
41
APEX 13(1-2) 1-61, 20 avr. 1998 Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
PLATE 2.
An example of occurrence of distinct, conspecific Oliva populations in neighbouring localities.
©. spicata (Rôding, 1798) in the vicinity of La Paz, Baja California, Mexico.
42
TURSCH, GREIFENEDER & HUART Oliva fulgurator and related species APEX 13(1-2): 1-61, 20 avr. 1998
©. rejecta
Balandra
SEA OF CORTEZ
DA Playa del Tesoro
Herendira
Bahia Pichilingue
Punta de Leon BAHIA Punta Prieta
Ensenada de
La Paz
@ collecting stations
43
APEX 13(1-2): 1-61, 20 avr. 1998
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
PLATE 3.
Some tips for quick identification.
=
TURSCH, GREIFENEDER & HUART Oliva fulgurator and related species APEX 13(1-2): 1-61, 20 avr. 1998
Figs. 1-2. Tip for quick identification of O. sayana (1). The angle of chevrons increases markedly during
growth, while in ©. fulgurator (2) the angle of chevrons stays nearly constant. Scale bars: 10 mm.
Fig. 3. Tip for quick identification of ©. scripta.
In most specimens, the angle of the fine
strokes pattern becomes progressively larger
as one goes towards the base, while the angle
of the dark chevrons remains nearly constant
8. O. sayana 9. ©. fulgurator 10. ©. scripta
Figs. 4-10. Tip for quick identification of O. polpasta (6): characteristic “cogwheel pattern” in apical view,
very rarely present in other related species. Note the very wide filament channel of O. scripta (10). Scale
bars: 10 mm.
45
APEX 13(1-2): 1-61, 20 avr. 1998 Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
PLATE 4. Type material
1-4. O. foxi Stingley, 1984.
1. HOLOTYPE ANSP 358356. (H: 36.98 mm, D: 14.82 mm). Cocos I., CosTA RICA.
2. PARATYPE ANSP 358357. (H: 37.60 mm, D: 15.17 mm). Cocos I., CosTA RICA.
3. PARATYPE AMNH 264800. (H: 34.9 mm, D: 14.1 mm). Cocos Is., CosTaA RICA.
4. PARATYPE AMS C.170890. (H: 33.35 mm, D: 13.36 mm). Cocos Is., CosTA RIcA. .
5. O. spicata deynzerae Petuch & Sargent, 1986.
5. HOLOTYPE USNM 841452 (H: 53.19 mm, D: 21.95 mm). Cocos Is., COSTA RICA.
6-9. O. polpasta Duclos, 1833.
6. O. polpasta Duclos, 1833. FIGURED SYNTYPE, MNHN (H: 40.1 mm; D: 21.0 mm).
7. O. truncata Marrat, 1867. HOLOTYPE, MCM (H: 37 mm, D: 18 mm).
8. O. kerstitchi da Motta, 1985. HOLOTYPE MHNG 984.631 (H: 24.5, D: 11.2 mm).
9. O. olssoni Petuch & Sargent, 1986. Holotype USNM 841444 (H: 35.03 mm; D: 19.57 mm). 60 m off Gulf of
Panama (8°7'N, 78°40'W), PANAMA.
46
TURSCH, GREIFENEDER & HUART Oliva fulgurator and related species APEX 13(1-2): 1-61, 20 avr. 1998
O. spicata
deynzerae
©. foxi ©. foxi
par. ANSP par. AMNH
©. polpasta ©. truncata ©. kerstitchi ©. olssoni
47
APEX 13(1-2): 1-61, 20 avr. 1998 Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
PLATE 5. Type material.
1-4. O. sayana Ravenel, 1834.
1. O. litterata Lamarck (not Rôding, 1798), 1811. Encycl., PI. 362, Figs. 1a, 1b.
2. O. sayana texana Petuch & Sargent, 1986. HoLoTYPE USNM 841465 (H: 43.60 mm; D: 17.54 mm). Shallow
water, Padre |., off South Texas.
3. ©. sayana sarasotensis Petuch & Sargent, 1986. HOLOTYPE USNM 841450 (H: 40.60 mm; D: 17.90 mm). 15 m,
sand, SCUBA, 15 km West of Lido Beach, Sarasota, Florida.
4. O. circinata citrina Johnson, 1911. HoLoOTYPE MCZ 6267 (H: 60.51 mm, D: 23.56 mm). “W. Coast Florida”.
5-10. ©. scripta Lamarck, 1811.
5. ©. scripta Lamarck (not Rôding, 1798), 1811. Encycl., PI. 362, Figs. 4a, 4b.
6. O. caribaeensis Dall & Simpson, 1901. Holotype USNM 159692 (H: 31.61 mm, D: 14.93 mm). Mayaguez, Puerto
Rico.
7. O. trujilloi Clench, 1938. HoOLOTYPE MCZ 57240 (H: 40.52 mm, D: 17.95 mm). "Puerto Plata, Santo Domingo,
Dominican Republic. Dredged 30 ft."
8. ©. trujilloi Clench, 1938. PARATYPE AMNH 79010 (H: 34.8 mm; D: 15.5 mm). "Dredged in 5 fath., Puerto Plata,
Santo Domingo".
9. ©. trujilloi Clench, 1938. PARATYPE (out of five) AMS C.095147, (H: 39.57 mm, D: 18.59 mm). "Suction dredge, 5
m, bluish clay, Puerto Plata, Santo Domingo”. Purchased Krause coll., 1972".
10. ©. scripta venezuelana Petuch & Sargent, 1986. HoLoTYPE USNM 841451 (H: 44.22 mm, D: 22.02 mm). 30 m
off Punta Fijo, Paranagua Peninsula, Venezuela.
48
TURSCH, GREIFENEDER & HUART
©. litterata ©. sayana
texana
#
©. sayana ©. sayana
sarasotensis citrina
NN /
d
DE
"4
Oliva fulgurator and related species
©. scripta
©. trujilloi
par. AMNH
APEx 13(1-2): 1-61, 20 avr. 1998
©. caribaeensis ©. trujilloi
©. trujilloi ©. scripta
par. AMS venezuelana
49
APEX 13(1-2): 1-61, 20 avr. 1998 Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
PLATE 6. Type material.
1-10. O. fulgurator (Rôding, 1798).
1. O. obesina Duclos, 1840. FIGURED SYNTYPE MNHN (H: 44.0 mm, D: 26.3 mm).
2. O. aldinia Duclos, 1845. FIGURED SYNTYPE MNHN (H: 55.6 mm, D: 30.3 mm).
3. O. porcea Marrat, 1870. HOLOTYPE MCM (H: 42 mm, D: 20.8 mm).
4. O. graphica Marrat, 1870. HOLOTYPE MCM (H: 44.6 mm, D: 22.2 mm).
5. ©. fusiformis Lamarck, 1811. Encycl., PI. 367, Figs. 1a, 1b.
6. O. jamaicensis Marrat, 1867. HOLOTYPE MCM (H: 37.7 mm, D: 18.4 mm).
7. O. oblonga Marrat, 1867. HOLOTYPE MCM (H: 63.2 mm, D: 29.2 mm).
8. O. magdae Petuch & Sargent, 1986. HOLOTYPE USNM 841441 (H: 37.57 mm, D: 18.62 mm). In fish trap, 300 m,
off Matanzas Bay, Cuba. C. J. Finlay, coll., 1956.
9. O. mercatoria Marrat, 1871. SYNTYPE MCM (H: 41.0 mm, D: 19.5 mm).
10. ©. fulgurator (Rôding, 1798). Martini Fig. 562, rotated.
50
TURSCH, GREIFENEDER & HUART Oliva fulgurator and related species APEX 13(1-2): 1-61, 20 avr. 1998
. obesina ©. aldinia ©. graphica ©. fusiformis
©. fulgurator
©. jamaicensis ©. oblonga ©. mercatoria
Si
APEX 13(1-2): 1-61, 20 avr. 1998 Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
PLATE 7. Type material.
1-9. ©. fulgurator (Rôding, 1798) (continued).
1. O. timoria Duclos, 1840. Figured SYNTYPE (out of 3) (H: 62.0 mm, D: 33.1 mm), MNHN.
2. O. jamaicensis zombia Petuch & Sargent, 1986. HOLOTYPE USNM 841454. (H: 23.87 mm, D: 12.61 mm). On
beach after storm, near sand flats, Southern side of Gonave |. Haiti. 1974.
. ©. broderipi Ducros, 1857. HOLOTYPE MNHN (H:27.8 mm; D: 13.4 mm).
. ©. contoyensis Petuch, 1988. HOLOTYPE USNM 859945 (H: 36.71 mm, D: 18.42 mm). 35 m. off Contoy |.
Yucatan, Mexico.
. ©. sargenti Petuch, 1987. Holotype USNM 859864 (H: 28.60 mm, D: 14.03 mm). 5 m, sand, Malmok, Aruba.
. ©. bifasciata jenseni Petuch & Sargent, 1986. Holotype USNM 859302 (changed from 841453, Germon in Jitt..)
(H: 50.30 mm, D: 24.12 mm). 0.5 m sand, off Blue Horizons Beach, Bermuda, R. Jensen coll. 1963.
. ©. drangai Schwengel, 1951. HOLOTYPE ANSP 247107. (H: 40.47 mm, D: 18.62 mm). Pigeon Pt, Tobago.
. ©. reticularis bollingi Clench, 1934. HoLoTYPEe MCZ 76656 (H: 61.95 mm, D: 28.52 mm). “Crab-trap, Miami,
Florida”.
9. O. nivosa Marrat, 1871. HOLOTYPE MCM (H: 49.2 mm, D: 22.0 mm).
10. O. bewleyi Marrat, 1870. HoLOTYPE MCM (H: 41.8, D: 18.1 mm).
&
oo on
œ |
52
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
©. timoria 2
©. bifasciata
jenseni
©. jamaicensis ©. broderipi
zombia
©. drangai ©. reticularis
bollingi
©. contoyensis ©. sargenti
O. nivosa ©. bewleyi
5
APEX 13(1-2} 1-61, 20 avr. 1998 Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
PLATE 8. Type material.
1-10. O. fulgurator (Rôding, 1798) (continued).
1. ©. maya Petuch & Sargent, 1986. HoLoTYPE USNM 859301 (changed from 841453, Germon jn litt.) (H: 57.81
mm: D: 24.97 mm). Trawled by shrimper in 35 m. off Contoy Is, Yucatan, Mexico. 1977.
2. O. goajira Petuch & Sargent, 1986. HOLOTYPE USNM 841433 (H: 36.79 mm, D: 15.49 mm). 60 m off Cabo La
Vela, Colombia.
. ©. figura Marrat, 1870. Holotype MCM (H: 41.2 mm, D: 18.1 mm).
4. O. bifasciata Küster in Weinkauff, 1878. FIGURED SPECIMEN SMF 9353. (H: 61.39 mm, D: 26.73 mm). “Brasilien”.
. ©. reticularis greenwayae Clench, 1937. HoLoTyYPE MCZ 115455 (H: 59.65 mm, D: 25.18 mm). “Smith Point,
Grand Bahama Island, Bahamas”.
6. ©. reticularis pattersoni Clench, 1945. HoLoOTYPE MCZ 151166 (H: 62.57 mm, D: 28.29 mm). “5 miles E of
Crabbing Point, Great Bahama Island, Bahamas”.
7. O. ernesti Petuch 1990. Holotype USNM 860535. (H: 33.66 mm, D: 15.45 mm) (not H: 38 mm as in description).
Trawled 40m silty sand, off Portobello, Panama (East).
8. O. finlayi Petuch & Sargent, 1986. HOLOTYPE USNM 841432 (H: 39.98 mm; D: 18.71 mm). 200 m, Matanzas Bay,
Cuba.
9. ©. reticularis Lamarck, 1811. Encycl!. PI. 361, Figs. 1a, 1b.
10. ©. bahamasensis Petuch & Sargent, 1986. Holotype USNM 841426 (H: 44.58 mm, D: 20.62 mm). In lobster
pot, in 200 m off Grand Bahama |., Bahamas.
(es)
an
54
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEX 13(1-2): 1-61, 20 avr. 1998
©. figura
©. pattersoni ©. ernesti ©. finlayi
©. bifasciata ©. reticularis
greenwayae
©. reticularis ©. bahamensis
55
APEX 13(1-2): 1-61, 20 avr. 1998 Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
PLATE 9. Type material.
1-10. O. fulgurator (Rôding, 1798) (continued).
1. O. antillensis Petuch & Sargent, 1986. HOLOTYPE USNM n° 841425 (H: 28.05 mm, D: 11.47 mm). On sand flats,
low tide, at night, Southern coast of Gonave Island, Haiti.
2. O. olorinella Duclos, 1835. SYNTYPE MNHN (H: 28.7 mm; D: 12.1 mm.).
. ©. bullata Marrat, 1871. HOLOTYPE MCM (H: 60.83 mm; D: 26.21 mm).
. ©. barbadensis Petuch & Sargent, 1986. HoLOTYPE USNM 841427 (H: 47.05 mm; H: 20.32 mm). 200 m off St.
James, Barbados.
5. O. bifasciata sunderlandi Petuch, 1987. HoLoTYPE USNM 859904 (H: 22.02 mm, D: 9.56 mm). Trawled 150 m
West of Cedar Key, Florida.
6. O. formosa Marrat, 1870. HOLOTYPE MCM (H: 45.23 mm; D: 19.52 mm).
7. O. reclusa Marrat, 1871. HOLOTYPE MCM (H: 43.2 mm, D: 18.1 mm).
8. O. circinata Marrat, 1871. HOLOTYPE MCM (H: 55.4, D: 23.4 mm).
9
. ©. circinata tostesi Petuch, 1987. HOLOTYPE USNM 859865 (H: 46.20 mm, D: 21.37 mm) (not H: 41 mm as in
description). 1 m, sand, llha Cabra, lllhabela, Sao Paulo State, Brazil.
10. ©. cribraria Marrat, 1883? HOLOTYPE MCM (H: 48.22 mm, D: 22.12 mm).
& ©
56
TURSCH, GREIFENEDER & HUART
Oliva fulgurator and related species
APEx 13(1-2): 1-61, 20 avr. 1998
©. antillensis
©. formosa ©. reclusa
©. bullata
©. circinata
JS 7
©. bifasciata
sunderlandi
©. circinata ©. cribraria
tostesi
sh
APEX 13(1-2): 1-61, 20 avr. 1998 Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
PLATE 10. Type material.
1-10. O. spicata (Rôding, 1798).
à
. ©. timoria Duclos, 1840. Figured SYNTYPE (out of 3) (H: 55.8 mm, D: 39.1 mm), MNAN.
N
. ©. pindarina Duclos, 1840. FIGURED SYNTYPE MNHN (H: 53.4mm; D: 27.3 mm).
[e)
. ©. ligneola Reeve, 1850. SYNTYPE (out of 2) BM(NH) 1892.9.24.4-5 (H: 25.60 mm, D: 13.09 mm). "Loc?
Purchased of Miss Steere".
H
. ©. punctata Marrat, 1870: SYNTYPE (out of two) MCM (H: 35.35 mm; D: 17.24 mm).
5. O. spicata (Rôding, 1798). Martini's figures 509 and 510 (rotated, cast shadow removed for better legibility).
. ©. radix Petuch & Sargent, 1986. HOLOTYPE USNM 841446 (H: 36.62mm, D: 22.51 mm). 75m, off Isla la Plata,
Ecuador.
oo
7. O. subangulata corteziana Petuch & Sargent, 1986. HoLoTyYPE USNM 841457 (H: 32.33 mm; D: 16.21 mm).
Northern end ofthe Gulf of California.
8. O. spicata hemphilli Johnson, 1911. SYNTYPE (out of five) ANSP 111697 (H: 54.05 mm, D: 23.38 mm). “San
Ignacio Lagoon, Lower California. John Ford colin.”
9. O. rejecta Burch & Burch, 1962. PARATYPE MNHN (H: 31.3, D: 13.6 mm). La Paz, Baja California, Mexico.
10. ©. rejecta Burch & Burch, 1962. PARATYPE USNM 667317 (H: 37.24 mm; D: 16.01 mm). La Paz, Baja California,
Mexico.
58
APEx 13(1-2): 1-61, 20 avr. 1998
Oliva fulgurator and related species
TURSCH, GREIFENEDER & HUART
©. spicata
©. pindarina ©. ligneola ©. punctata
©. timoria 1
©. rejecta
©. rejecta
par. MNHN
©. spicata
©. subangulata
©. radix
par. USNM
var. hemphilli
corteziana
59
APEX 13(1-2) 1-61, 20 avr. 1998 Oliva fulgurator and related species TURSCH, GREIFENEDER & HUART
PLATE 11. Type material.
1-8. O. spicata (Rôding, 1798) (continued).
1. O. cumingii Reeve, 1850. SYNTYPE (out of 3) BM(NH) 1987008 (H: 48.4 mm, D: 24.2 mm). "Gulf of California. H.
Cuming colin."
2. O. fuscata Marrat, 1870. SYNTYPE (out of 3) MCM (H: 42.66 mm; D: 20.30 mm).
3. O. brunnea Marrat, 1870. HOLOTYPE MCM (H: 34,0 mm, D: 16.1 mm).
4. O. oniska Duclos, 1844. FIGURED SYNTYPE MNHN (H: 61.0 mm; D: 26.2 mm).
5
. ©. spicata perfecta Johnson, 1911. SYNTYPE (out of three) ANSP 111729 (H: 59.06 mm, D: 25.73 mm). “West
Coast Central America. John Ford colin.”
. ©. subangulata Philippi, 1848. SYNTYPE (out of two) BM(NH) 1924.1.5.98-99. (H: 59.62 mm, D: 26.38 mm).
. ©. fuscata Marrat, 1870. Another SYNTYPE (out of 3) MCM (H: 39.76 mm; D: 18.62 mm).
. ©. violacea Marrat, 1867.
. ©. intertincta Carpenter, 1857. SYNTYPE (largest of two) USNM 716187, (both specimens numbered with ink
"592/2121") (H: 19.68 mm, D: 9.70 mm). Mazatlan, Sinaloa, Mexico, ex coll. F. REIGEN.
10. O. ionopsis Berry, 1969. Holotype SMBNH 34656 (H: 30.54 mm, D: 13.76 mm). 33fms, Bahia de las Palmas,
Baja California.
© © 1
60
TURSCH, GREIFENEDER & HUART Oliva fulgurator and related species APEX 13(1-2): 1-61, 20 avr. 1998
©. brunnea
©. cumingi ©. fuscata 1 ©. oniska ©. spicata var.
perfecta
©. subangulata ©. fuscata 2 ©. violacea ©. intertincta ©. ionopsis
61
tas leve 6 fe htr es L
: en
Dee
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Lo
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D Cou oo ta fée TATÉ 0-5 MMA
PACAUD
Nouveau genre de Pholadidae
Un nouveau genre de Pholadidae (Mollusca, Bivalvia) du
Paléogène inférieur du nord de l'Atlantique
Jean-Michel PACAUD
5, Avenue Pablo Picasso - F - 94120 Fontenay-sous-Bois, France
MOTS CLES. Pholadidae, Paléogène inférieur, Atlantique, Europe, Etats Unis, Systématique.
KEY WORDS. Pholadidae, Early Paleogene, Atlantic Ocean, Europe, United States, Systematics.
RESUME. La position systématique de toutes les espèces de la sous-famille des Pholadinae du
Paléogène inférieur est revue. Les caractères morphologiques propres à ces espèces permettent de les
séparer du genre Barnea où elles étaient jusqu'ici classées. La comparaison avec les autres genres de
Pholadinae conduit à proposer un nouveau genre, Cyrtopleuropsis. Ces espèces sont seulement
connues en Atlantique et sont les suivantes: pour l'Europe, Cyrtopleuropsis orbignyana (Lévesque in
Graves, 1847), C. freyteti (Plaziat, 1970), C. lescailloni nov. sp. et pour les Etats-Unis, C. alatoidea
(Aldrich, 1886). Une distribution stratigraphique de l'ensemble des Pholadinae cénozoïques est
donnée.
ABSTRACT. The aim of this work is a review of the systematics of all Pholadinae from the Lower
Paleogene. Except Cyrtopleuropsis lescailloni wich is new, these species are currently assigned in the
genus Barnea. However comparisons with Barnea and other genera of Pholadinae show distinctive
morphological characters wich permit to propose a new genus, Cyrtopleuropsis. Paleogene species are
only know an Atlantic Ocean: for the Europe,Cyrtopleuropsis orbignyana (Lévesque in Graves,
1847), C. freyteti (Plaziat, 1970), C. lescailloni nov. sp. and for the United States, C. alatoidea
APEX 13(1-2): 63-75, 20 avr. 1998
(Aldrich, 1886). Stratigraphic distribution is given for all caenozoic Pholadinae.
INTRODUCTION
L’Examen des Pholadinae du Paléogène inférieur du
Nord de l’Atlantique m'avais conduit à rapporter
Pholas orbignyana (Lévesque in Graves, 1847), de
l’Eocène du Bassin Anglo-parisien, dans le genre
Cyrtopleura Tryon, 1862 (LE RENARD & PACAUD,
1995). De nouvelles récoltes de ce matériel rare et
généralement mal conservé permettent des
observations supplémentaires pour préciser la position
systématique de ce groupe de Pholadidae, aboutissant à
proposer un nouveau genre: Cyrtopleuropsis.
Abbréviations utilisées
MNEAN-LP: Laboratoire de Paléontologie. Muséum
National d'Histoire Naturelle de Paris.
MNAN-BIMM: Laboratoire de Biologie des
Invertébrés Marins et Malacologie. Muséum National
d'Histoire Naturelle de Paris.
UPMC: Service de la collection de Paléontologie,
Université Pierre et Marie Curie, Paris VI.
BMNH: The Natural History Museum, Londres.
USNM: National Museum of Washington.
PRI: Paleontological Research Institution, Ithaca, New
York.
ETUDE SYSTEMATIQUE
Famille: PHOLADIDAE Lamarck, 1809
Sous-Famille: Pholadinae Lamarck, 1809
Genre: Cyrtopleuropsis nov. gen.
Espèce-type: Pholas orbignyana Lévesque in Graves,
1847 (= Pholas levesquei Watelet, 1851)
Derivatio nominis. Forme fossile rappelant par sa
morphologie le genre Cyrtopleura. Genre grammatical:
féminin.
Diagnose. Coquille allongée. Bord antérieur acuminé.
Bord postérieur atténué. Bord antéro-ventral échancré
et sinueux. Bord cardinal retourné sur le crochet.
Réflexion umbonale non cloisonnée. Charnière
pourvue d'un chondrophore. Apophyse large et
arrondie. Surface dorsale présentant une sculpture
63
APEX 13(1-2): 63-75, 20 avr. 1998
Nouveau genre de Pholadidae
PACAUD
radiaire s'interrompant à mi-longueur de la valve,
découpant en festons le bord antérieur. Région dorso-
postérieure marquée par de nombreuses granulations.
Distribution. La présence de Pholades dans le Bassin
de Paris fut relevée pour la première fois dans le
Cuisien par Lévesque (1847) avec Pholas orbignyana.
L'espèce fut découverte plus tard par l'abbé LAMBERT
dans les sables sparnaciens de Sinceny (Aisne). D'après
un moule interne provenant de la collection Armand
BAZIN, DESHAYES (1857, p. 135) signale P. orbignyana
du Thanétien de Gannes (Oise). PERREAU (WYNS & al.
1981, p. 50) cite avec doute cette espèce du Thanétien
de Bachivillers (Oise) d'après une valve incomplète
(matériel non conservé, comm. Perreau). DESHAYES
(1857, p. 135) signale aussi P. orbignyana au Lutétien
d'après des fragments provenant de Grignon (Yvelines)
et un exemplaire récolté par DUTEMPLE à Damery
(Marne). L'espèce est également signalée par de
nombreux auteurs dans l'Eocène d'Angleterre. Au
Paléocène des Etats-Unis, une autre espèce est décrite
par ALDRICH (1886) de Gregg's Landing (Alabama),
Pholas alatoidea. PLAZIAT (1970) a décrit une espèce
du Sparnacien des Corbières septentrionales, Barnea
freyteti, proche de Pholas orbignyana.
Discussion générique. La fragilité du test des Pholades
rend très rare la conservation complète à l'état fossile
de leurs valves et de leurs plaques accessoires
(protoplax, mesoplax et metaplax). Après la mort de
l'animal, ces plaques accessoires sont dispersées et
généralement détruites. Leurs formes énigmatiques
furent à l'origine de méprises de la part de divers
auteurs. (COSSMANN (1907, p. 201-202) décrit
l'apophyse de Cyrtopleuropsis orbignyana comme
Avicula moloti du Sparnacien de Pourcy (Marne). Des
protoplax ou des apophyses furent aussi déterminés
comme Scutum, Crepidula, Acmaea ou Patella
(TURNER, 1954). Ces plaques accessoires permettent
une distinction générique. Les protoplax de Barnea
Leach in Risso, 1826 à sillon médian prononcé et de
Anchomasa Leach, 1852 sans sillon médian se
distinguent de celui de Monothyra Tryon, 1862 plus
triangulaire et de celui de Cyrtopleura Tryon, 1862, en
forme de T. LAURIAT-RAGE & MAKINSKY (1983) ont
signalé la présence de plusieurs protoplax d'une
pholade dans le Miocène moyen de Thenay (Indre-et-
Loire) et ont ainsi pu rapporter au sous-genre
Monothyra, l'espèce Pholas dujardini Mayer, 1859.
GLIBERT & VAN DE POEL (1966, p. 14-15) plaçaient
alors cette dernière dans le sous-genre Thovana Gray,
1947.
Aucune de ces plaques n'a été retrouvée dans le
Bassin de Paris. De ce fait l'interprétation générique
fondée sur le protoplax ne peut être établie et repose
donc sur la forme de la coquille, de l'apophyse et sur la
présence à la charnière d'un chondrophore,
modification de la charnière servant de surface
d'insertion au résilium.
64
Les genres Pholas (s. str.) Linné, 1758 (espèce-type
P. (s. str.) dactylus Linné, 1758), Pholas (Thovana)
Gray, 1847 (espèce-type P. (Thovana) campechiensis
Gmelin, 1791), et Pholas (Monothyra) Tryon, 1862
(espèce-type P. (Monothyra) orientalis Gmelin, 1791)
(Figs. 23-28) se caractérisent par le cloisonnement
alvéolaire de 1a réflexion umbonale et diffèrent donc
des coquilles paléogènes. Chez l'espèce-type de Barnea
(s. str.) Leach in Risso, 1826, B. (5. str.) candida
(Linné, 1758), l'ornementation radiaire atteint
l'extrémité postérieure, l'échancrure antéro-ventrale est
à peine indiquée et l'extrémité antérieure est arrondie,
non baillante, l'apophyse est d'une forme étroite et
courbe. Un chondrophore est présent chez B. candida,
cependant il est nettement plus faible et moins projeté
que celui des coquilles paléogènes (Figs. 29-30).
L'échancrure antéro-ventrale des coquilles paléogènes
rappelle celle de Zirfaea Leach in Gray, 1847 (espèce-
type Zirfaea crispata (Linné, 1758) (Figs. 33-34), mais
la région médiane des valves de Zirfaea montre un
sulcus umbono-ventral,correspondant à la clavicule de
la face interne, ce qui exclut tout rapprochement.
L'espèce-type du sous-genre Barnea (Anchomasa)
Leach, 1852, B. (Anchomasa) parva (Pennant, 1777)
est plus proche par son échancrure antéro-ventrale
sinueuse, par son extrémité antérieure en forme de
rostre ainsi que par ses côtes radiaires s'interrompant à
mi-longueur de la valve (Figs. 31-32). Toutefois la
forme des valves, tordue, courte et large, souvent
tronquée dans la région postérieure, l'échancrure
ventrale plus longue (elle atteint la moitié de la
longueur totale), la forme de son apophyse, étroite et
fortement courbée, écartent tout autre rapprochement
avec les coquilles paléogènes.
C'est finalement avec les coquilles de Cyrtopleura (s.
str.) Tryon, 1862 et Cyrtopleura (Scobinopholas) Grant
& Gale, 1931 que Cyrtopleuropsis s'apparente le plus
(tableau 1). L'ornementation des coquilles paléogènes
concorde assez bien avec celle de l'espèce-type de
Cyrtopleura (s. str.), C. (s. str.) cruciger (Sowerby,
1834). Toutefois les valves de cette dernière présentent
une sculpture radiaire se poursuivant au-delà de la
région médiane, s'atténuant progressivement au lieu de
cesser brusquement vers le milieu de la longueur (Figs.
19-20). L'apophyse est d'une forme étroite, fortement
courbée et creusée, alors que celles observées encore
en place sur les charnières de Cyrtopleuropsis
orbignyana (Lévesque in (Graves, 1847), de C.
alatoidea (Aldrich, 1886) et de C. lescailloni sont plus
petites, plus larges et plus arrondies. Les fortes côtes
radiaires de la région antérieure de Cyrtopleuropsis
rappellent celles que l'on observe chez l'espèce-type du
sous-genre Scobinopholas Grant & Gale, 1931,
Cyrtopleura (Scobinopholas) costata (Linné, 1758);
toutefois chez Scobinopholas la sculpture radiaire
persiste sur toute la surface dorsale des valves et le
bord antérieur est nettement arrondi (Figs. 21-22),
l'apophyse est large et arrondie. Les empreintes
musculaires nettes ainsi que le sinus palléal large et
profond de Cyrtopleura (s. str.) rappelle assez celui de
PACAUD
Nouveau genre de Pholadidae
APEX 13(1-2): 63-75, 20 avr. 1998
Cyrtopleuropsis. Chez Scobinopholas le sinus palléal
est invisible et certaines coquilles de Cyrtopleura
(Scobinopholas) costata montrent, en dessous de
l'empreinte de l'adducteur postérieur, une saillie
donnant plus d'assise pour l'attache du muscle. La
présence chez Cyrtopleuropsis d'un chondrophore,
visible seulement chez quelques Pholades, notamment
chez Cyrtopleura (s. str.), la rapproche du groupe
Cyrtopleura-Scobinopholas. Il faut cependant
remarquer l'absence de granulations sur la surface
dorso-postérieure et d'échancrure ventrale chez
Cyrtopleura et Scobinopholas. L'extrémité antérieure,
arrondie et à peine baillante, contraste avec les
coquilles paléogènes. TRACEY (communication
personnelle) observe que C. orbignyana ne se récolte
que dans des gisements à substrat non induré. C
[P Plaques accessoires accessoires inconnues! + |
Présence ee chondrophore
Cyrtopleu-| Cyrto- | Scobino- | Pholas onothyra| Thovana | Barnea lnchomas | Zirfaea
ropsis pleura pholas
Présence de granulations sur la ; :
face dorsale
orbignyana se rencontre dans des gisements tels que
Abbecourt, Bachivillers, Cuise-la-Motte, Pierrefonds,
Trosly-Breuil, Liancourt-Saint-Pierre, Le Rocquet et
Sinceny où le substrat semble avoir été sableux ou
sablo-vaseux. Ceci va dans le sens de notre
rapprochement générique car les Cyrtopleura actuelles
sont inféodées aux substrats meubles, tandis que
Barnea, genre dans lequel les auteurs classaient
Jusqu'ici les diverses espèces discutées ici, creuse des
niches dans la roche. Une morphologie et une écologie
comparable suggèrent des relations étroites entre
Cyrtopleuropsis et les genres Cyrtopleura et
Scobinopholas qui en dérivent probablement (d’où le
rapprochement fait par TRACEY (1986), LE RENARD &
PACAUD (1995) et JEFFERY & TRACEY (1997) pour
Pholas orbignyana).
PRES OR ES RER RS ES
PR RS CS RE ER
LS RCE RE TR RE ES RS CS RS EE
Sinus es large et Su pal age apré |
a mpreintes
arquées
Présence d'une échancrure antéro-
entrale
Présence d'un sulcus umbono-
ompant à mi-longueur de la
alve
Ornementation
Tableau 1.
Cyrtopleuropsis orbignyana
(Lévesque in Graves, 1847) nov. comb.
(Figs, 1-9 et 13)
1847 Pholas Orbignyana Lévesque in Graves : 643.
Synonymie:
1851 Pholas Levesquei Watelet : 11, pl. I, fig. 1-5.
1906 Avicula? Moloti Cossmann & Pissarro : pl. XLV,
fig. 121-14.
1907 Avicula Moloti Cossmann & Pissarro : 201-202,
pl. VIL fig. 121-14.
1963 Barnea cf. levesquei (Watelet) -
pl. 8, fig. 1-5; pl. 9, fig. 1-2.
1963 Barnea cingulata Stinton : 69, pl. 8, fig. 6a-b.
1981 Barnea sp. Perreau in Wyns & al. : 50
Stinton : 68-69,
Te CP RC
Ornementation radiaire couvrant
la totalité de la surface dorsale
radiaire
s'attenuant au-delà de la mi-
longueur de la valve
| invisible |
Autres références:
1850 Pholas orbignyana Lévesque
d'Orbigny : 321, n° 439.
1856 Pholas Levesquei Watelet - Deshayes : pl. VL fig.
10-12; p. 135-136 (1857).
1882 Pholas Levesquei Watelet - Zittel :
b.
1885 Pholas Orbignyi Lévesque in Graves - Chelot :
192:
1886 Pholas Levesquei Watelet - Fritel : pl. IL, fig. 9.
1886 Barnea Levesquei (Watelet) - Cossmann : 13.
1891 Dactylina Levesquei (Watelet) - Newton : 94.
1930 Barnea levesquei (Watelet) - Wrigley : 378.
1986 Cyrtopleura levesquei (Watelet) - Tracey : 118.
1995 Cyrtopleura orbignyana (Lévesque in Graves) -
Le Renard & Pacaud : 67.
in (Graves -
138, fig. 197a-
APEX 13(1-2): 63-75, 20 avr. 1998
1996 Cyrtopleura orbignyana (Lévesque in Graves) -
Pacaud & Le Renard : 184.
1996 Cyrtopleura levesquei (Watelet) - Tracey, Todd,
Le Renard, King & Goodchild : 136-137.
1997 Cyrtopleura levesquei (Watelet) - Jeffery &
Tracey : 89-90, pl. 6, fig. 15.
Localité-type. Cuise-la-Motte (Oise), Cuisien.
Distribution de l'espèce. Thanétien -
(Paléocène - Eocène).
Yprésien
Matériel-type. Non retrouvé.
Matériel thanétien. Sables de Bracheux, Bachivillers,
lieu-dit " Remise Madame " (Oise): 1 valve gauche,
Fig. 5 (coll. Pons), 1 fragment, valve droite (coll.
Pacaud, lot n° P25359); 1 fragment, valve droite (coll.
Lescaillon), Abbecourt, lieu-dit " Bois des Godins "
(Oise): 1 fragment (coll. Pacaud, lot n° P24889).
Autre matériel examiné. Cuisien: Cuise-la-Motte
(Oise), 1 valve droite, longueur: 32 mm., Figs. 1-2
(coll. Lhomme, MNHN-LP, lot n° B50646), 1 valve
gauche, longueur: 27 mm., Figs. 3-4 (coll. Lhomme,
MNEAN-LP, lot n° B50646), 1 valve droite mutilée,
Figs. 6-7 (coll. Pacaud, lot n° P32275), 1 valve gauche
incomplète, Figs. 8-9 (coll. Pacaud, lot n° P32275); 20
valves (coll. Marceaux, MNHN-LP); Pierrefonds
(Oise): 23 exemplaires en connection (coll. Munier-
Chalmas, UPMC), Trosly-Breuil (Oise): 1 valve droite
(coll. Pacaud, lot n° P36113);, Le Roquet (Oise): 2
valves gauches et 1 valve droite incomplètes (coll.
Pacaud, lot n° P36304). Sparnacien: Sinceny (Aisne): 2
valves droites incomplètes (coll. Pacaud, lot n°
P45360).
Description du matériel thanétien. Coquille de 97
mm. de long et de 29 mm. de large, fortement
inéquilatérale. Bord antérieur acuminé, terminé par un
rostre. Bord postérieur long et atténué. Bord antéro-
ventral échancré et sinueux. Bord cardinal fortement
Nouveau genre de Pholadidae
PACAUD
retourné, enveloppant le crochet. Réflection umbonale
sinueuse et non cloisonnée. Chondrophore bilobé, lame
inférieure courte. Apophyse large et arrondie. Surface
dorsale présentant toujours, dans la région antérieure,
une sculpture radiaire composée de 16 côtes fortement
marquées dans le test s'interrompant à mi-longueur de
la valve. Les huit premières sont puissantes, largement
espacées, gravant profondément leurs empreintes dans
la face interne de la valve; se comportant comme des
carènes lisses sur la face interne de la valve, se
projetant en pointes aiguës sur le bord antéro-ventral, le
découpant en festons plus ou moins profonds. Sur la
face externe, les côtes sont en revanche puissamment
sculptées de chevrons épineux imbriqués les uns dans
les autres. La sculpture radiaire qui suit est constituée
de 8 autres côtes plus rapprochées et cesse
brusquement vers le milieu de la longueur. Leur
ornementation est plus faible. La surface interne des
valves conserve également leurs traces sous la forme de
lignes martelées par les petites écailles externes. Elles
sont bien moins gravées dans le test que les huit
premières et leur inclinaison umbono-antéro-ventrale
est à l'opposé de ces dernières. La région postérieure
est parcourue de lignes de croissance sublamelleuses,
régulières, faiblement marquées et de très nombreuses
granulations rappelant celles qui peuvent s'observer à la
surface de Panopea Ménard ou de Poromya Forbes.
L'impression du muscle adducteur postérieur est ovale,
étroite et allongée. Le sinus palléal est large, aigu en sa
partie palléo-ventrale, remontant très haut vers
l'impression, petite et ovalaire, du muscle adducteur
antérieur.
Comparaison avec le matériel cuisien. La sculpture
de la face dorsale et l'inclinaison umbono-antéro-
ventrale des costules rayonnantes sont identiques
lorsque l'on compare la valve de la Figure 5 du
Thanétien et celles des des Figures 1- 4 du Cuisien de
Cuise-la-Motte. D'autre part, l'identité des valves du
Cuisien et de celles du Thanétien se retrouve sur les
traces musculaires, le chondrophore et l'apophyse. Le
test moins convexe des coquilles de Bachivillers est dû
à une déformation par compaction post-sédimentaire.
L'attribution du matériel thanétien à Cyrtopleuropsis
orbignyana ne fait aucun doute.
Figs. 1-2. Cyrtopleuropsis orbignyana (Lévesque in Graves, 1847), valve droite de Cuise-la-Motte (Oise), Eocène
inférieur (Cuisien), collection Lhomme, MNHN-LP, lot n° B50646. (Photo L. Merlette). x 2. Figs. 3-4.
Cyrtopleuropsis orbignyana (Lévesque in Graves, 1847), valve gauche de Cuise-la-Motte (Oise). Eocène inférieur
(Cuisien), collection Lhomme, MNHN-LP, lot n° B50646. (Photo L. Merlette). x 2. Fig. 5. Cyrtopleuropsis orbignyana
(Lévesque in Graves, 1847), valve gauche de Bachivillers (Oise). Paléocène (Thanétien supérieur), collection Pons.
(Photo P. Lozouet). grandeur nature, longueur. 97 mm., largeur: 29 mm. Figs. 6-7. Cyrtopleuropsis orbignyana
(Lévesque in Graves, 1847), valve droite de Cuise-la-Motte (Oise). Eocène inférieur (Cuisien), collection Pacaud.
ru: réflexion umbonale; cu: callosité umbonale; maa: empreinte du muscle adducteur antérieur, map: empreinte du
muscle adducteur postérieur; a: apophyse; c: chondrophore. (Photo P. Lozouet). x 2. Figs. 8-9. Cyrtopleuropsis
orbignyana (Lévesque in Graves, 1847), valve gauche de Cuise-la-Motte (Oise). Eocène inférieur (Cuisien),
collection Pacaud. c: chondrophore. (Photo P. Lozouet). grandeur nature. Figs. 10-11. Cyrtopleuropsis lescailloni
nov. sp., valve droite de Chamery (Marne). Eocène moyen (Lutétien moyen), holotype, MNHN-LP, lot n° R11583.
(Photo P. Lozouet). x 1,5. Fig. 12. Cyrtopleuropsis lescailloni nov. sp., valve gauche de Chamery (Marne). Eocène
moyen (Lutétien moyen), collection Pons. (Photo P. Lozouet). grandeur nature.
66
Nouveau genre de Pholadidae APEX 13(1-2): 63-75, 20 avr. 1998
maa
APEX 13(1-2}) 63-75, 20 avr. 1998
Barnea cingulata Sunton, 1962, (Figs. 13) fondée sur
un unique exemplaire (BMNAH, lot n° LL14708) du
Lutétien de Selsey (Balanophyllia Bed, Sussex,
Angleterre) ne diffère de C. orbignyana que par un
contour palléo-ventral nettement arrondi avant de se
prolonger par un bord postérieur atténué. Les dernières
côtes radiaires sont plus espacées dans la région
médiane, cependant, l'échancrure antéro-ventrale ainsi
que la sculpture radiaire sont analogues. JEFFERY &
TRACEY (1997) considèrent Barnea cingulata comme
synonyme de C. orbignyana, ce que je soutiens
également. Le contour particulier du bord palléo-
ventral n'est qu' un caractère individuel de l'holotype.
J'ajouterai que C. orbignyana est signalé au Paléocène
et à l'Eocène en Angleterre par NEWTON (1891),
WRIGLEY (1930), STINTON (1962), TRACEY (1986)
ainsi que TRACEY & al. (1996).
Cyrtopleuropsis lescailloni nov. sp.
(Fig. 10-12)
Synonymie:1857 Pholas Levesquei Deshayes partim,
non Watelet : 135 (non pl. 6, fig. 10-12).
Etage-type. Lutétien moyen (Eocène moyen).
Localité-type. Chamery, lieu-dit "Les
(Marne).
Beurges"
Derivatio nominis. Espèce dédiée à Michel Lescaillon.
Matériel-type. Holotype, Figs. 10-11 (MNHN-ELP, lot
n° R11583, récolte Lescaillon), valve droite.
Dimensions de l'holotype. Longueur: 75 mm., largeur:
18 mm.
Autre matériel Chamery, lieu-dit "Les Beurges"
(Marne): une valve droite (coll. Lescaillon); une valve
gauche, Fig. 12 (coll. Pons); une valve gauche (coll.
Boucher); Damery (Oise) fide DESHAYES; Grignon
(Yvelines) fide DESHAYES.
Diagnose. Coquille allongée, fortement inéquilatérale.
Nouveau genre de Pholadidae
PACAUD
Bord antérieur court, acuminé. Bord postérieur long et
atténué. Valve épaisse et convexe. Bord antéro-ventral
échancré et sinueux. Région antérieure de la surface
dorsale ornée d'une sculpture radiaire composée de 15
à 17 côtes, bien marquées dans le test, s'interrompant
au premier tiers de la longueur de la valve. Dans sa
partie extrême, les 9 ou 10 premières côtes sont
puissamment sculptées de chevrons imbriqués les uns
dans les autres. Leur courbure importante applique une
très nette convexité au bord antérieur. Elles sont
largement espacées. Ces espaces sont ornés par de
nombreuses lignes de croissance fines et serrées. Les
côtes gravent leurs empreintes dans la surface interne
sous forme de carènes lisses. Dans sa partie médiane, la
sculpture radiaire de la surface dorsale est constituée de
6 à 7 côtes dont l'espacement est moindre et dont
l'ornementation est plus faible que sur les côtes
antérieures, leur inclinaison est plus importante et elles
sont festonnées de petites écailles allongées. La surface
interne est à peine marquée par ces côtes. Région
postérieure de la surface dorsale parcourue par de
nombreuses lignes de croissance sublamelleuses,
régulières et bien marquées. De très nombreuses
granulations couvrent cette surface, essentiellement
dans la partie postérieure de la coquille.
L'impression du muscle adducteur postérieur, ovale,
étroite et allongée est située très haut. Le sinus palléal
est large, aigu en sa partie palléo-ventrale, il remonte
très haut vers l'impression, petite et ovalaire, du muscle
adducteur antérieur. Le bord cardinal est fortement
retourné et enveloppe le crochet. La réflexion
umbonale est non cloisonnée. La charnière est pourvue
d'un solide chondrophore, aigu à son extrémité.
L'apophyse est large et arrondie.
Comparaison. C. lescailloni se distingue de C.
orbignyana par:
- une sculpture radiaire moins vigoureuse. Les espaces
entre les costules sont nettement mieux festonnés par
les lignes de croissance. Les côtes les plus proches du
bord antérieur rejoignent la sinuosité de l'échancrure
antéro-ventrale, donnant une nette convexité au bord
antérieur (Fig. 12), tandis qu'elles sont fuyantes chez C.
orbignyana, où pratiquement toutes les costules
rejoignent le rostre antérieur (Fig. 8). L'ornementation
rayonnante occupe chez C. lescailloni le tiers de la
longueur totale, contre la moitié de la longueur chez C.
orbignyana..
Fig. 13. Barnea cingulata, valve droite de Selsey (Sussex, Angleterre). Eocène moyen (Lutétien), holotype, BMNH,
lot n° LL 14708. (d'après Stinton, 1963). x 2. Fig. 14. Cyrtopleuropsis freyteti (Plaziat, 1970), valve gauche de
Montlaur (Aude). Sparnacien. (d'après Plaziat, 1970). x 2,5. Fig. 15. Pholas aldrichi, fragment de valve gauche de
Bell's Landing (Alabama, Etats-Unis). Paléocène (Thanétien), holotype, USNM, lot n° 643735. (d'après de Gregorio,
1890). x 2. Figs. 16-17. Cyrtopleuropsis alatoidea (Aldrich, 1886), valve gauche de Gregg's Landing (Alabama,
Etats-Unis). Paléocène (Thanétien), syntypes, USNM, lot n° 638792. (d'après de Gregorio, 1890). x 2. Fig. 18.
Cyrtopleuropsis alatoidea (Aldrich, 1886), valve gauche de Gregg's Landing (Alabama, Etats Unis), Paléocène
(Thanétien), syntype, PRI, lot n° 183. (d'après Harris, 1897). x 2. Figs. 19-20. Espèce-type du sous-genre
Cyrtopleura (s.str.), C. (s.str.) cruciger (Sowerby, 1834). (d'après Turner, 1954). x 2. Figs. 21-22. Espèce-type du
sous-genre Cyrtopleura (Scobinopholas), C. (S.) costata (Linné, 1758). (d'après Turner, 1954). x 0,7.
68
PACAUD
Nouveau genre de Pholadidae
(A
C RZ
vs,
227
APEX 13(1-2): 63-75, 20 avr. 1998
69
APEX 13(1-2): 63-75, 20 avr. 1998
- une surface dorsale parcourue par de plus nombreuses
lignes de croissance. Elles festonnent les côtes
antérieures de chevrons moins importants et nettement
plus imbriqués les uns dans les autres. Le contour de
l'échancrure antéro-ventrale est de ce fait nettement
moins découpé que celui de C. orbignyana.
- un bord antérieur plus nettement arrondi, moins
acuminé. Le contour de la réflexion umbonale
enveloppant le crochet est arrondi tandis qu'il est
sinueux chez C. orbignyana.
- un chondrophore plus fort, plus nettement bilobé sur
la valve gauche. La lame inférieure est plus longue,
plus aigüe.
- une face interne moins marquée par la sculpture
radiaire où seules les côtes les plus proches du bord
antérieur sont visibles. Les stries de croissance
sublamelleuses ne se distinguent absolument pas à
l'intérieur de la coquille comme on peut le voir chez C.
orbignyana.
- une position plus haute et plus étroitement allongée
de l'impression du muscle adducteur postérieur, placée
pratiquement contre le bord palléal. La pointe aiguë du
sinus palléal est plus étroite.
Cyrtopleuropsis freyteti
(Plaziat, 1970) nov. comb.
(Fig. 15)
1970 Barnea freyteti Plaziat : 61-62, fig. 41; pl. IX, fig.
15a-b.
Matériel-type. Syntypes, SE de Montlaur, Aude (coll.
Plaziat, lots n° 1836 et n°1838); SW de Montlaur,
Aude (coll. Plaziat, lot n° 1839), NW de Villemagne,
Aude (coll. Plaziat, lot n° 1840).
Distribution. Yprésien (Sparnacien, Eocène inférieur).
Dimensions de l'exemplaire figuré (Syntype).
Longueur: 34 mm.; largeur: 11 mm.
Observations. (Cette espèce, contemporaine de
Cyrtopleuropsis orbignyana (Sables sparnaciens de
Sinceny, Aisne) s'en écarte toutefois par un
allongement moins important, par un nombre de côtes
moindre (12), et par l'inclinaison plus faible des côtes
postérieures. L'ornementation rayonnante n'occupe
seulement que le tiers de la longueur totale. Les
coquilles ne sont pas isolées mais soudées à des
plaquettes de calcaires bioclastiques et ne montrent
pour la plupart que leur face externe. Les lots n° 1839
Nouveau genre de Pholadidae
PACAUD
et 1840 montrent cependant leur face interne mais la
charnière nous est inconnue et je ne peux donc discuter
ni du chondrophore ni de l'apophyse; cependant les
caractères morphologiques de la face dorsale de cette
espèce, la sinuosité du bord antéro-ventral ainsi que la
projection du bord antérieur, confirment l'appartenance
de Barnea freyteti au genre Cyrtopleuropsis.
Cyrtopleuropsis alatoidea
(Aldrich, 1886) nov. comb.
(Fig. 15-18)
1886 Pholas alatoidea Aldrich : 36, pl. 4, fig. 9a-c.
Synonymie.
1848 Pholas Roperiana Tuomey : 153. nomen nudum.
1890 Pholas Aldrichi de Gregonio : 237, pl. 38, fig. 17.
1893 Barnea alatoidea Aldrichi (de Gregorio) -
Cossmann : 5.
1965 Pholas sp. Palmer & Brann : 266.
Autres références.
1890 Pholas alatoidea Aldrich - de Gregorio : 237, pl.
38, fig. 15-16.
1893 Barnea alatoidea (Aldrich) - Cossmann : 5.
1897 Pholas alatoidea Aldrich - Harris : 69, pl. 13, fig.
15-15a
1899 Pholas alatoideus Aldrich - Harris & Veatch :
304, pl. 53, fig. 12.
1960 Pholas alatoidea Aldrich - Brann & Kent : 686.
1965 Pholas alatoidea Aldrich - Palmer & Brann : 265.
Matériel-type. Syntypes, Gregg's Landing, Alabama
(Etats Unis): 2 valves gauches, dont une incomplète
(USNM, lot n° 638792), Bell's Landing, Alabama
(Etats Unis): 1 valve gauche incomplète (holotype du
Pholas aldrichi, USNM, lot n° 643735).
Autre matériel Gregg's Landing, Alabama (Etats
Unis): 1 valve gauche et 1 valve droite incomplète
(PRI, lots n° 183 et 184), Sabinetown, Texas (Etats
Unis): " Pholas sp. " matériel perdu.
Distribution. Thanétien (Tuscahoma Formation et
Sabinetown Formation, Paléocène).
Dimensions. Valve gauche syntype (Fig. 16)
Longueur: 50 mm; largeur: 14 mm.
Figs. 23-24. Espèce-type du sous-genre Pholas (s.str), P. (s.str) dactylus Linné, 1758. Port-la-Nouvelle, Aude
(coll. Pacaud). grandeur nature. Figs. 25-26. Espèce-type du sous-genre Pholas (Monothyra), P. (M.) orientalis
Gmelin, 1791. (d'après Turner in Moore, 1969). x 1. Figs. 27-28. Espèce-type du sous-genre Pholas (Thovana), P.
(T.) campechiensis Gmelin, 1791. (d'après Turner, 1954). x 0,5. Figs. 29-30. Espèce-type du sous-genre Barnea
(s.str.), B. (s.str.) candida (Linné, 1758). Port-la-Nouvelle, Aude (coll. Pacaud). x 2. Figs. 31-32. Espèce-type du
sous-genre Barnea (Anchomasa), B. (A.) parva (Pennant, 1777). (d'après Turner, 1954). x 1. Figs. 33-34. Espèce-
type du genre Zirfaea, Z. crispata (Linné, 1758). (d'après Turner, 1954). x 1,5.
70
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= LT
AT
ET IE SRI RE
71
APEX 13(1-2): 63-75, 20 avr. 1998
Nouveau genre de Pholadidae
PACAUD
Observations. Pholas alatoidea Aldrich, 1886 du
Thanétien de Gregg's Landing (Alabama, Etats Unis)
appartient comme C. orbignyana, C. freyteti et C.
lescailloni, au genre Cyrtopleuropsis par ses caractères
morphologiques, par la forme de son apophyse et par la
présence à la charnière d'un chondrophore. En outre,
HARRIS (1897) figure une autre valve gauche (Fig. 18)
provenant de la même localité, et sur laquelle on
distingue nettement de nombreuses granulations
couvrant la partie postérieure de la coquille.
Cyrtopleuropsis alatoidea diffère de C. orbignyana par
son bord antérieur moins acuminé, sans échancure
antéro-ventrale nette. Les lignes de croissance
festonnent les côtes antérieures de petites écailles, le
contour antéro-ventral est de ce fait nettement moins
découpé que celui de C. orbignyana L'espèce
américaine se rapproche bien plus de C. lescailloni par
son extrémité antérieure non rostrée, par son
échancrure antéro-ventrale peu marquée. Mais
l'orientation des côtes les plus proches du bord
antérieur est différente: projetées vers l'extrémité
antérieure chez C. alatoidea, les côtes rejoignent la
sinuosité de l'échancrure antéro-ventrale chez C.
lescailloni. L'ornementation rayonnante occupe
seulement le tiers de la longueur totale chez C.
lescailloni et la moitié chez C. alatoidea. Une des
figures de DE (GREGORIO (1890) montre un
chondrophore étroit remontant vers la callosité
umbonale (Fig. 17), alors qu'il est projeté vers
l'intérieur de la coquille chez C. orbignyana et chez C.
lescailloni. Pholas aldrichi de Gregorio, 1890 (Fig. 15)
du Thanétien de Bell's Landing (Alabama, Etats-Unis),
fondé sur un fragment de la région antérieure, est tout à
fait semblable à C. alatoidea.
Pour toutes ces raisons, je considère C. alatoidea
comme une espèce distincte des espèces du Bassin
Anglo-parisien. Elle est toutefois contemporaine de C.
orbignyana; les unités lithostratigraphiques
américaines "Bells Landing Marl Member" et "Greggs
Landing Marl Member" , de la Tuscahoma Formation,
sont actuellement attribuées au Thanétien (SIESSER &
al., 1985).
CONCLUSIONS
Cyrtopleuropsis orbignyana, C. freyteti, C. lescailloni
et C. alatoidea ne sont connues qu'au Paléogène
inférieur du Nord de l'Atlantique. Elles ont en commun
un bord antérieur fortement ornementé, un rostre
antérieur bien individualisé, une sculpture radiaire
s'interrompant dans la région médiane des valves, des
granulations sur la surface dorso-postérieure, une
apophyse arrondie et surtout une échancrure antéro-
ventrale sinueuse. Les espèces actuelles et fossiles de
Cyrtopleura et de Scobinopholas appartiennent à un
groupe très voisin qui diffère par l'absence d'une
échancrure antéro-ventrale nette, par une atténuation
progressive de l'ornementation rayonnante qui se
poursuit au-delà du milieu de la longueur et par la
forme de l'apophyse, plus étirée chez Cyrtopleura (voir
72
tableau 1).
D'après mes observations, l'existence de Barnea et de
Pholas (Monothyra) dès le Crétacé (TURNER, 1954;
TURNER in MOORE, 1969) doit être remise en question.
Je n'ai pas en effet pu observer la présence de ces deux
genres dès le Mesozoïque.
STEPHENSON (1952) a décrit une espèce du
Cénomanien (Crétacé sup.) de Cook County et de
Grayson County de la "Woodbine Formation" (Texas,
Etats Unis), Pholas? scaphoides. Je ne connais cette
espèce que d'après les figures des faces externes qu'en
donna l'auteur, ainsi que la figure reproduisant
l’holotype donnée par KENNEDY (1993). On ne peut en
préciser le statut générique, ne distinguant ni un
éventuel cloisonnement alvéolaire de la réflexion
umbonale (brisée?) ni l'intérieur des valves. KENNEDY
(1993, p. 398) classe Pholas scaphoides, ainsi qu’une
nouvelle espèce du Coniacien et du Santonien (Crétacé
sup.) du Nord de la Californie, dans le genre Barnea
(Anchomasa). Son choix est motivé uniquement sur
l’apparente absence de cloisonnement alvéolaire de la
réflexion umbonale. KELLY (1988), dans son inventaire
des Pholades mésozoïques, considère P. scaphoides
comme le plus ancien Pholadinae et rapporte cette
espèce au genre Pholas (Monothyra). Les faces
externes de Pholas scaphoides et de Barnea
(Anchomasa) saulae Kennedy, 1993 montrent une
ornementation radiaire dépassant la mi-longueur des
valves ainsi qu'une très nette échancrure antéro-
ventrale. DARTEVELLE & FREINEX (1957) rapportèrent à
Pholas scaphoides un moule externe de petite
dimension et un fragment plus grand de moule interne
provenant des niveaux crétacés du Congo (Afrique).
Ces deux espèces, dont les caractères morphologiques
des faces externes sont assez comparables à ceux de
Pholas (s. str.) et de Cyrtopleuropsis, restent de
classement douteux. Elles appartiennent
vraisemblablement au même genre, toutefois le
rattachement au genre Barnea (Anchomasa) n’est pas
convaincant.
Pholas pectorosa Conrad, 1852, espèce-type du
genre Clavipholas Conrad, 1868, a été décrit du
Crétacé supérieur des Etats Unis (STEPHENSON, 1941).
Toutefois, la possible fermeture du bord antérieur par
un callum(TURNER in MOORE, 1969) chez Clavipholas
pectorosa semble jeter un doute sur l'assimiliation de
ce genre à la sous-famille des Pholadinae et serai
probablement plus à sa place parmi les Martesiinae. On
pourra ainsi noter les affinités de cette espèce avec
Pholas petrosa Conrad, 1842. L'exemplaire figuré par
CLARK (1896) de l'Eocène du Maryland (Etats Unis)
montre un galbe et une ornementation tout à fait
semblable à Clavipholas pectorosa. PALMER & BRANN
(1965) rapportent Pholas petrosa au genre Phenacomya
Dall, 1898. Ce genre est actuellement considéré comme
synonyme du Martesiinae Eutylus Vincent, 1891. Ceci
me conduit, dans l'ignorance des caractères internes des
valves de l'espèce du Crétacé, au vu de sa morphologie
et de ses affinités, à envisager de rattacher cette
dernière plutôt aux Martesiinae.
PACAUD
Nouveau genre de Pholadidae
APEX 13(1-2): 63-75, 20 avr. 1998
Il ressort donc que Cyrfopleuropsis est le plus ancien
Pholadinae décrit du Cénozoïque (tableau 2). Pholas
scaphoides Stephenson, 1953 et Barnea (Anchomasa)
saulae Kennedy, 1993 sont les seules Pholadinae
mésozoïques connus. À ma connaissance, hormis
Zirlona increnata (Marwick, 1929) du "Duntroonian"
(Oligocène supérieur, voir BEU & MAXWELL, 1990) de
Nouvelle Zélande, aucun autre Pholadinae n'a été décrit
de l'Oligocène. La présence de Cyrtopleuropsis
alatoidea (Aldrich, 1886) dans le Paléocène de
l'Alabama et du Texas marque ainsi le premier jalon
d'un groupe qui, sur les côtes américaines, évoluera
vers celui de Cyrfopleura-Scobinopholas. Au Miocène,
Cyrtopleura (Scobinopholas) arcuata (Conrad, 1841)
est décrite de Nansemond River près de Sufifolk en
Virginie. CAMPBELL (1993) la signale également dans
les sédiments miocènes de Yorktown, de Smithfield et
de Chuckatuck en Virginie. DALL (1898) et HANNA
(1926) signalent la présence de C. (Scobinopholas)
costata (Linné, 1758) au Pliocène de Caloosahatchie
marls en Floride et à Coyote Mountain en Californie.
DALL (1898) signale C. (S.) costata au Pléistocène dans
le Massachusetts, dans le Maryland et en Floride.
Actuellement, l'aire de distribution de Cyrfopleura (5.
str.) est restreinte aux côtes du Mexique, du Costa Rica
et du Panama. Celle de C. (Scobinopholas) est plus
étendue; elle comprend, sans discontinuité, les côtes du
Massachussets jusqu'aux côtes du Texas. C.
(Scobinopholas) est également signalé sur les côtes
brésilienne, uruguayenne et argentine.
Le tableau 2 montre donc la répartition
stratigraphique des genres de la sous-famille des
Pholadinae, l'echelle chronostratigraphique utilisées est
celle proposée par ODIN & ODIN (1990).
REMERCIEMENTS. Je tiens à remercier Steve Tracey
et Jean-Claude Plaziat pour les informations qu'ils
m'ont apportées durant l'élabortion de cette note:
Michel Lescaillon, Jacques Pons et Pascal Boucher
pour le prêt de leur matériel; Didier Merle et le Pr.
Agnès Lauriat-Rage pour la relecture du manuscrit. Je
remercie également le Pr. Lauriat-Rage qui m'a permis
de consulter les collections du Laboratoire de
Paléontologie (MNHN-LP), Mme Denise Pajaud pour
m'avoir laissé consulter la collection Munier-Chalmas
(UPMC), ainsi que Mme Suzanne Freneix (MNHN-
LP) pour les informations sur les faunes crétacées
qu'elle m'a apportées. Je remercie aussi Pierre Lozouet
(MNEAN-BIMM) ainsi que Lionel Merlette (MNHN-
LP) pour les ciichés photographiques.
REFERENCES
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73
APEX 13(1-2} 63-75, 20 avr. 1998 Nouveau genre de Pholadidae PACAUD
EE
ACTUEL
- 0,01 MA
PLEISTOCENE
- 1,6 MA
PLIOCENE
- 5,3 MA
MIOCENE
- 23,5 MA
=
CA
=
Ÿ
mi
L_ 4
ra
4
_
_
ln
L 4
- 34 MA
EOCENE
- 53 MA
MARTESIINAE ?
PALEOCENE
- 65 MA
CRETACE
Tableau 2. Distribution stratigraphique de Cyrtopleuropsis et des genres voisins de Pholadinae: 1, Zirfaea Leach in
Gray, 1842; 2, Barnea (Anchomasa) Leach, 1852; 3, Pholas (s.str.) Linné, 1758; 4, Pholas (Thovana) Gray, 1847;
5, Pholas (Monothyra) Tryon, 1862; 6, Cyrtopleura (Scobinopholas) Grant & Gale, 1931; 7, Cyrtopleura (s.str.)
Tryon, 1862; 8, Barnea (s.str.) Leach in Risso, 1826; 9, Cyrtopleuropsis nov. gen.; 10, Pholas an Cyrtopleuropsis ?
(?Pholas scaphoides Stephenson, 1952); 11, Clavipholas Conrad, 1868; 12, Zirlona Finlay, 1930.
74
PACAUD
Nouveau genre de Pholadidae
APEX 13(1-2): 63-75, 20 avr. 1998
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76
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PACAUD
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TUOMEY, M. 1848. Report on the Geology of south
Carolina, in 4°, Columbia : 293 pp.
TURNER, RD. 1954. The family Pholadidae in the
Western Atlantic and Eastern Pacific. I: Pholadinae.
Johnsonia 3(33): 1-64.
TURNER, R.D. in MOORE, R.C. 1969. Treatise on
Invertebrate Paleontology, Part. N, Mollusca 6,
Bivalvia, Pholadacea. The Geological Society of
America, Inc., The University of Texas : pp. N702-
N741, figs. E162-E213.
VINCENT, E.G. 1891. Contribution à la paléontologie
de l'Eocène Belge. Pholadidae. Annales de la Société
royale Malacologique de Belgique 26: 164-168.
WATELET, À. 1851. Recherches faites dans les sables
tertiaires inférieurs des environs de Soissons. Bulletin
de la Société Archéologique, Historique et
Scientifique de Soissons 5: 113-126.
WRIGLEY, A. 1930. Notes on English Eocene boring
Mollusca, with descriptions of new species.
Proceedings of the Geologists' Association of London
40(4): 376-383.
ZITTEL K..A. 1882. Handbuch der Palaeontologie.
Palaezoologie. 11. Mollusca und Arthropoda. R.
Oldenbourg, München und Leipzig : 1109 pp.
VAN BELLE & DELL’ ANGELO
New species of Ischnochitonidae
APEX 13(1-2): 77-79, 20 avr. 1998
Description of a new species Zschnochiton dolii sp. nov.
(Polyplacophora: Ischnochitonidae) from Civitavecchia, Italy
Richard A. VAN BELLE’ and Bruno DELL'ANGELO?
! Scientific Collaborator, Koninklijk Belgisch Instituut voor Natuurwetenschappen,
Vautierstraat 29, B-1000 Brussels, Belgium
? Via Mugellese 66D, 59100 Prato, Italy
KEY WORDS. Polyplacophora, Ischnochitonidae, Zschnochiton dolii sp. nov., Italy.
ABSTRACT. /schnochiton dolii sp. nov. is described. It is compared with all recent Mediterranean
and northeastern Atlantic Zschnochiton species, and with the European fossil species from the
Pliocene and the Miocene.
RESUME. /schnochiton dolii sp. nov. est décrit et comparé à tous les Zschnochiton récents de la
Méditerranée et du nord-est de l’Océan Atlantique, et aux espèces fossiles européennes du Pliocène et
du Miocène.
INTRODUCTION
On June 1991, a Roman jar (=dolium ; IV century
B.C.) was found in the Tyrrhenian Sea, off
Civitavecchia, at a depth of 550 meters (Tringali &
Ardovini, 1993). In the muddy contents 333 valves (69
head, 228 intermediate and 36 tail) were found of a
possibly subfossil Polyplacophora species which
proved to be new to science.
SYSTEMATICS
Family ISCHNOCHITONIDAE Dail, 1889
Subfamily Ischnochitoninae
Genus Zschnochiton Gray, 1847.
Type species :
1847.
Chiton textilis Gray, 1828, SD Gray,
Ischnochiton dolii sp. nov.
Figs 1-5
Type material. Off Civitavecchia, -550 m, 333 valves
: holotype, 3 x 7.5 mm (Fig. 1) + 5 paratypes in Museo
di Zoologia of the Bologna University (holotype reg.
no 11302, paratypes reg. no 11303), other paratypes in
Institut Royal des Sciences Naturelles de Belgique (4
valves, reg. no IG 28523), Museum National d'Histoire
Naturelle, Paris (3 valves), R. Ardovini coll. (138
valves), F. Giovini coll. (80 valves), B. Dell’ Angelo
coll. (90 valves, reg. no P151F/01), R.A. Van Belle
coll. (12 valves, reg. no F1003àa).
Tuscan Archipelago, -70/100 m : 1 intermediate
valve, 2.2 x 5.5 mm (B. Dell’ Angelo coll. reg. no
L15/01).
Type locality. Tyrrhenian Sea, off Civitavecchia, Italy,
-550 m.
Diagnosis. Dimensions of largest valves : head valve
3.5 x 7 mm. intermediate valve 3 x 7.5 mm, tail valve 4
x 6.5 mm. Considering the size of these valves and the
fact that all Zschnochiton species are oval to elongate
oval, one can assume that Z. dolii was an animal of
medium size (up to about 20 mm long), rather elevated
(jugal angle 85-110°), carinated with the side slopes
straight, the valves not beaked. Colour of tegmentum
dark brown, variously blotched and/or streaked with
white.
Description. Head valve (Fig. 2) semicircular, front
slope straight to very weakly concave, posterior margin
widely V-shaped, deeply notched in the middle,
tegmentum sculptured with rather flat, subgranulose
radiating ribs, 17-26 near apex, splitting up to more
than double that number near periphery of valve,
17
APEX 13(1-2): 77-79, 20 avr. 1998
interstices very narrow, ribs concentrically crossed by
numerous growth lines. Valve II pentagonal, other
intermediate valves (Figs 1, 4) broadly rectangular,
front margin obtusely angular, straight and slightly
slanting at both sides of jugal angle, side margins
weakly convex, hind margin straight, mucro
inconspicuous, lateral areas moderately elevated,
sculptured like head valve, 4-5 radiating ribs, in some
valves becoming obsolete towards side margin, central
area with 13-19 longitudinal sulci (Fig. 5) on both sides
of the smooth, wedge-shaped jugum, the 3-4 innermost
sulci forwardly converging towards jugum, intervening
nbs rather flat, slightly wider than sulci. Tail valve
(Fig. 3) depressed, short, about twice as wide as long,
front margin broadly angular like in intermediate
valves, hind margin less than semicircular, mucro
shghtly antemedian, little raised, antemucronal area
sculptured like central area of intermediate valves,
postmucronal area with 18-20 radiating, flattish ribs,
crossed by rather deep concentric grooves, in some
valves this sculpture becomes indistinct near the
mucro.
Articulamentum whitish, apophyses wide, short,
evenly rounded, jugal sinus shallow, relatively narrow,
weakly convex, insertion plates short, slit formula 9-
10/1/7-8 (one intermediate valve has a second slit on
one side), slits inequidistant, upper area of teeth
grooved, eaves very narrow.
Distribution. Only known from the type material.
Remarks. As the subgeneric division of Zschnochiton
s.l. is partly based on characteristics of the perinotum
(here lacking), a subgeneric assignment for Z. dolii is
impossible.
A similar remark equally applies to all fossil
Ischnochiton species.
Etymology. The specific name refers to the Roman jar
(= dolium) in which the valves have been found.
Discussion. Comparing the recent European
Ischnochiton species with Z. dolii, we conclude that the
latter differs from them all.
1. (Stenosemus) albus (Linnaeus, 1767), an Atlantic
species, has the tegmentum of all valves uniformly
microgranulose, smooth and glossy to the naked eye.
Apart from the tegmental colour and the always
present, wavy concentric lirae on end valves and lateral
areas of intermediate valves, Z (Simplischnochiton)
rissoi (Payraudeau, 1826), common in the
Mediterranean Sea, has a much greater number of
longitudinal riblets on the central areas.
1. (Stenosemus) exaratus (G.O. Sars, 1878), an
Atlantic deep water species, differs by the uniformly
dirty white to yellowish tegmental colour, the weaker
sculpture and especially, on the central areas, the
78
New species of Ischnochitonidae
VAN BELLE & DELL’ ANGELO
longitudinal sulci which do not reach the posterior
margin of the valve.
IL. (Simplischnochiton) obtusus Carpenter in Pilsbry,
1893, described from Portugal, has the lateral areas
more strongly raised, only 10 longitudinal sulci on the
pleural areas, and the mucro of the tail valve median
and prominent.
I. dolii seems closest related to Z (Stenosemus)
vanbellei Kaas, 1985, known from the white coral
banks of the Ligurian Sea, but that species has only 16
non splitting, radiating nbs on the head valve, 2 ribs (3-
4 in valve IT) on the lateral areas of the intermediate
valves, and only 7-9 longitudinal sulci on either side of
the jugum.
As the correct geological age of Z. dolii cannot be
established with any degree of certainty, Z. dolii needs
to be compared with European Pliocene and Miocene
Ischnochiton species too.
Both /. anserinus Laghi, 1977, from the Pliocene of
the Modena Basin, Italy, and Z. ulivii Dell’ Angelo &
Forli, 1996, from the Pliocene of Pietrafitta, Italy, have
a completely different sculpture, without longitudinal
or radiating rbs.
1. rudolticensis Sulc, 1936, from the Miocene of the
Vienna Basin, Austria, has a sculpture resembling that
of Z. (S.) rissoi and consequently cannot be conspecific
with Z. dolii.
I. korytnicensis Baluk, 1971, from the Miocene of the
Holy (Cross Mountains, Poland, has undulating
longitudinal ribs on central area of intermediate valves,
and irregular nodular elevations, variable in outline, on
lateral areas and end valves.
ACKNOWLEDGEMENTS.
We are indebted to R. Ardovini (Roma), M. Pizzini
(Roma) and F. Giovine (Villa S. Giovanni) who
provided the material, to C. Lombardi (Museo di
Zoologia, Bologna University) for preparing the SEM
photos, and E. Ulivi (Firenze) for the photo of the
holotype.
REFERENCES
BALUK, W. 1971. Lower Tortonian chitons from he
Korytnica clays, southern slopes of the Holy Cross
Mts. Acta geologica polonica 21 : 449-471.
DELL’ ANGELO, B. & M. FORLI. 1996. Two new
Polyplacophora species from theTuscan Pliocene. La
Conchiglia 28 (279), Suppl. : 42-49.
KAAS, P. & R.A. VAN BELLE. 1990. Monograph of
Living Chitons (Mollusca : Polyplacophora). Vol. 4,
Suborder Ischnochitonina : Ischnochitonidae :
Ischnochitoninae (continued). Additions to Vols 1, 2
and 3. E.J. Brill ed., Leiden, 298 pp.
LAGHI, G.F. 1977. Polyplacophora (Mollusca)
neogenici dell’ Appennino settentrionale. Boll. Soc.
paleontol. ital. 16 : 87-115.
VAN BELLE & DELL’ ANGELO New species of Ischnochitonidae APEX 13(1-2): 77-79, 20 avr. 1998
SULC, J. 1936. Studie über die fossilen Chitonen. I. Die TRINGALI, L. & R. ARDOVINI. 1993. Breve nota sul
fossilen Chitonen im Neogen des Wiener Beckens rivenimento di Odostomia silesui Nofroni, 1988 dalle
und der angrenzenden Gebiete. Ann. naturh. Mus. acque laziali (Mollusca : Gastropoda :
Wien 47 : 1-31. Heterobranchia). Notiz. CISMA 14 : 47-48.
Figs. 1-5. /schnochiton dolii sp. nov. Fig. 1. Holotype, 3 x 7.5 mm, Bologna Museum 11302 (photo E. Ulivi). Figs.
2-5. Paratypes, Bologna Museum 11303 (SEM photos C. Lombardi). Fig. 2. Head valve. Fig. 3. Tail valve. Fig. 4.
Intermediate valve. Fig. 5. Detail of sculpture of pleural area of intermediate valve. Scales - 2-4 ; 1 mm; 5: 100 pm.
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SOMMAIRE
C. Olabarria The influence of physicochemical parameters on the distribution
V. Urgorri of dominant bivalve species in the ensenada do Baño
J. S. Troncoso (Ria de Ferrol) in Northwest of Spain. 81.
R. Houart Description of eight new species of Muricidae (Gastropoda). 95:
J. Vidal Taxonomic revision of the Indo-Pacific Yasticardium assimile
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R. Houart Description of Trophon iarae n. sp., a new muricid from southern
West Atlantic (Gastropoda: Muricidae) with illustration
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OLABARRIA & al.
Bivalves and relationships with environmental parameters
APEX 13(3): 81-93, 20 juil. 1998
The influence of physicochemical parameters on the distribution
of dominant bivalve species in the ensenada do Baño
(Ria de Ferrol) in Northwest of Spain
C. OLABARRIA , V. URGORRI ? & J. S. TRONCOSO
! Fac. Ciencas del Mar, Univ. Auténoma de Sinaloa, C/Paseo Claussen s/n, 82000 Mazatlän (Sinaloa), Mexico
? Depart. Bioloxia Animal, Fac. de Bioloxia. Universidade de Santiago, E-15706, Santiago (La Coruña), Spain
* Depart. de Recursos Naturais e Medio Ambiente, Fac. de Ciencias do Mar, Universidade de Vigo, Vigo, Spain
KEY WORDS. Bivalves, synecology, physicochemical parameters, Ria de Ferrol, Spain.
MOTS CLEFS. Bivalves, synécologie, facteurs physico-chimiques, Ria de Ferrol, Espagne.
ABSTRACT. This paper presents a synecological study of the dominant bivalve species in the
Ensenada do Baño (Ria de Ferrol, NW Spain), and reports on relationships between the distribution of
these species and physicochemical parameters (particle size, carbonate content, organic matter
content, sorting coefficient and depth/ height with respect to tidal zero). We conclude that the most
important factors governing species distribution are grain size and depth/height.
RÉSUMÉ. Une étude synécologique des bivalves dominants de l' Anse do Baño (Ria de Ferrol, NO
Espagne) est présentée. Cette étude montre les relations entre leur distribution et les conditions
physico-chimiques du milieu (granulométnie, teneur en carbonates et en matière organique, coefficient
de sélection et profondeur). Cette étude a permis de mettre en évidence que les facteurs déterminant
de la distribution des bivalves sont le gradient sédimentaire et la profondeur ou la hauteur par rapport
au niveau 0 de la marée.
INTRODUCTION
In early studies on the biology of benthic faunas, an
understanding of sedimentological parameters proved
to be of utmost importance, due to their effect on the
faunal composition of communities and to the close
relationship between sediment variables and the
ecological preferences of the different species. This 1s
particularly true for infaunal species, which the nature
of the sediment determines life style as well as trophic
and reproductive habits.
In a number of studies, the primary goal has been to
gather information on relationships between mollusc
distributions and the physicochemical properties of the
sediment. It has been observed that the relationship
between mollusc distributions and the sedimentary
environment is the result of a dynamic interdependence
among physical, chemical and mucrobiological
sediment factors (BADER, 1954). Important studies in
this field include those of LANDE (1975), TUNBERG
(1981) and CORNET (1985, 1986) on the distribution
and ecology of bivalve communities of European
Atlantic coast, and those of DRISCOLL & BRANDON
(1973), FRANZ (1976), FRANZ & MERRILL (1980) on
the American Atlantic coast in which benthic
molluscan assemblages in relation to sediment were
studied. Previous studies about distribution and
ecology of molluscs of the coasts of northern Spain
include those of FIGUERAS (1956), VIÉITEZ (1976),
PLANAS & MORA (1984), LABORDA & MAZÉ (1987),
MAZÉ & LABORDA (1988), BORJA (1988, 1991) and
TRONCOSO & URGORRI (1992).
Despite of abundance of malacological studies on the
Galician coasts, there is a scarcity of research on
sinecology of molluscs. For that, we decided to extend
the knowledge of this group, particularly in the Ria de
Ferrol, where this group has been poorly studied. In the
present study, we investigated relationships between
bivalve distributions and physicochemical factors in the
Ria de Ferrol. The study was carried out in the
Ensenada de Baño (Baño Inlet), which contains a wide
range of sediment types (including mud, muddy sand,
shell gravel, coarse sand and maërl).
METHODS
The study area is located on the southern side of the
Ria de Ferrol (NW Spain), between Punta do Faro da
Palma (43°27'52"N; 08°16'49"W) and Punta Piteira
(43°27'57"N; 08°15'37"W), and has an area of 0.5 km?
and a maximum depth of 18 m (Fig. 1).
81
APEX 13(3): 81-93, 20 juil. 1998
Bivalves and relationships With environmental parameters
OLABARRIA & al.
The inlet is oriented in direction NNE-SSW; the
prevailing winds are southwesterlies for most of the
year, except in summer when northeasterlies become
dominant. The mean tidal range in the ria is 2,7 m, and
tidal effects give rise to strong currents (up to 1,5 m/s
in the ria’s central channel). Outward movement of
water from the ra provokes movement to the SE within
the inlet, while movement into the ria provokes
movement to the SSW within the inlet. These currents.
which are stronger at the mouth of the inlet than at
more distal points, are the dominant factor affecting
sediment distribution within the inlet.
Mollusc were sampled between July 1991 and June
1992 at 40 intertidal and 35 subtidal stations (one
sample per station). Sampling points were selected
along 12 parallel transects drawn across the inlet at
100 m intervals, taking samples at the points that were
Judged by visual examination to show a change in
nature, texture or substrate covering. In the subtidal
zone, the samples were collected by scuba diving. At
each point, a 0,25 m? square sample was taken, to a
depth of approximately 20 cm, using a rectangular
shovel. The tidal range at intertidal stations was
2.29 m. The intertidal samples were additionally
collected at the ends of each transect and, in the inner
intertidal zone, samples were also taken every 100 m
along each transect (OLABARRIA ef al. 1996). AI
samples were subsequently wet-sieved through a series
of sieves with 10, 2, and 0.5 mm mesh. Finally the
sieved samples were transported to the laboratory to be
sorted by the remounting technique (ROS, 1975).
Surface sediment samples were also collected from
each sampling point, for granulometric analysis and for
determination of organic matter and carbonate content
by the method of GUITIAN & CARBALLAS (1976)
(Table 1).
For each species and each environmental factor,
possible relationships between population density
(number of individuals in the sample) and the level of
that factor were investigated by Spearman rank
correlation analysis (SOKAL & ROHLF, 1979) (Table 2).
RESULTS
Distribution of individual species with respect
to sediment characteristics
The samples vielded a total of 7579 specimens of
bivalves belonging to 52 species, of which 11 (Myrilus
edulis, Thyasira flexuosa, Mysella bidentata,
Papillicardium papillosum, Parvicardium exiguum,
Cerastoderma edule, Abra alba, Venus verrucosa,
Dosinia exoleta, Venerupis senegalensis and Hiatella
arctica) represented 80.8% of the total, with Mysella
bidentata being the most representative species
(35.6%) (Table 3).
Mytilus edulis occurred most commonly in the
intertidal zone (93.3%). of individuals. versus 6.7% in
the subtidal zone reaching its highest densities in
coarse sand bottoms located in the intertidal zone on
the border of the inlet (Fig. 2A). The density of
82
individuals in the intertidal zone was positively
correlated with gravel, coarse sand, and carbonate
contents, and negatively correlated with fine sand
content (Table 2).
Thyasira flexuosa was found almost exclusively in
the subtidal zone (99,4% of individuals), reaching its
highest dénsities in the eastern area of the inlet (Fig.
2B), on fine sand bottoms with silt-clay contents of
over 10% and organic matter contents of 1-2%. There
Was a strong positive correlation between the density of
this species in the subtidal zone and fine sand and
organic matter contents, and a strong negative
correlation with coarse sand and carbonate contents
(Table 2).
Mysella bidentata was the most abundant and widely
distributed bivalve, with a broad distribution
throughout the inlet and a large number of individuals
in the subtidal zone (Fig. 2C). Correlation analysis did
not reveal any relationship with physicochemical
parameters in the subtidal zone, due to the widespread
occurrence of this species on all types of bottom. In the
intertidal zone, however, the density of this species
correlated positively with coarse sand and gravel
contents, and negatively with fine sand content (Table
2);
Papillicardium papillosum occurred most commonly
in the subtidal zone (96.7% of individuals). The highest
densities were those observed in the outer subtidal zone
(Fig. 2D), dominated by coarse sand and medium sand
bottoms with organic matter contents of less than 1% in
most cases. The density of this species in the subtidal
zone was positively correlated with coarse sand content
and depth, and negatively correlated with fine sand
content (Table 2).
Parvicardium exiguum Was commonest in the
intertidal zone (86.4% of individuals). The highest
densities were found in the mid-intertidal zone where
there 1s à Zostera noltii meadow (Fig. 3A). The density
of this species in the subtidal zone showed a significant
though weak positive correlation with medium and fine
sand contents, and a significant negative correlation
with gravel and carbonate contents. In the intertidal
zone, the density of this species was negatively
correlated with tidal height (Table 2).
Cerastoderma edule was present only in the intertidal
zone, and reached its highest densities in the inner
intertidal zone (Fig. 3B), where the bottoms are mostly
medium and fine sand with organic matter content
ranging between 0.5 and 1%. Correlation analysis did
not reveal significant relationships with the
physicochemical parameters, except for a negative
correlation with tidal height (Table 2).
Abra alba was most frequent in the subtidal zone
(845% of individuals) However, it was widely
distributed in the inlet (Fig. 3C), being most common
on bottoms with particle size smaller than 0.5 mm and
organic matter content greater than 1%. The density of
this species in the subtidal zone correlated positively
with the sorting coefficient and less strongly with
organic matter content, and negatively with coarse sand
OLABARRIA & al. Bivalves and relationships with environmental parameters
APEX 13(3): 81-93, 20 juil. 1998
Ria de FERROL
#
:
43°28"N
Ensenada |
8°14'W
do Baño
D 43°28'N
Punta S115
Redonda or 16
ST 510 s113 S114 o
da Palma °$102 o cote Punta Pitcira
EL
1101
rx
Mugardos
8°16 W
Fig. 1. Location of the study area and the sampling sites. A: Spain, B: Galicia, C: Ria de Ferrol, D: Ensenada do
Baño.
83
APEX 13(3} 81-93, 20 juil. 1998
Bivalves and relationships with environmental parameters
OLABARRIA & al.
content. In the intertidal zone, density showed a weak
but significant positive correlation with silt-clay
content (Table 2)
Venus verrucosa occurred largely in the subtidal
zone (945% of individuals), and reached its highest
densities on bottoms with coarse sediments and varying
percentages of organic matter (Fig. 3D). The density of
this species in the subtidal zone was negatively
correlated with fine sand content. In the intertidal zone,
however, a positive correlation was observed between
the density of this species and both coarse sand content
and the sorting coefficient (Table 2).
Dosinia exoleta occurred most commonly in the
subtidal zone (96.8% of individuals), and reached its
Lu Jen [Sc TS: Jos Jon Joux Eau Jos [sc Ts Jo Jon Jon
highest densities in the outer part of the inlet on coarse
sand bottoms with variable organic matter content (0.4-
1.7%) (Fig. 4A). The density of this species correlated
positively with coarse sand content and to a lesser
extent with carbonate content, and negatively with fine
sand content (Table 2).
Venerupis senegalensis was slightly more frequent in
the subtidal zone (61.8% of individuals), attaining its
highest densities in the outer inlet. In the intertidal zone
it was distributed homogeneously throughout the whole
area. In both cases, it occurred on bottoms of varied
physicochemical characteristics (Fig. 4B). In the
intertidal zone, density was negatively correlated with
tidal height (Table 2).
Table 1. Characteristics of the sampling sites. Qso : median particle size (mm), S-C : silt and clay contents (% w/w);
So : Sorting coefficient, CA : carbonate content (% w/w), OM : organic matter content (% w/w); D : depth (m) with
respect to tidal zero (subtidal sites), H : height (m) with respect to tidal zero (intertidal sites).
84
OLABARRIA & al. Bivalves and relationships with environmental parameters APEX 13(3): 81-93, 20 juil. 1998
16
.4 ; "924 1096,60 *
.142 180
L] a
152.156 #4 *
ee +
Fig. 2. Estimated densities (individuals per m2) of Mytilus edulis (A), Thyasira flexuosa (B), Mysella bidentata (C)
and Papillicardium papillosum (D) at sites at which that species was found
85
APEX 13(3) 81-93, 20 juil. 1998
Hiatella arctica was likewise slightiv more frequent
in the subtidal zone (655% of individuals), largely
occurring on bottoms with à particle size greater than
0.5 mm, both in the subtidal and intertidal zone (Fig.
4C). The density of this species in the subtidal zone
showed a slight positive correlation with gravel content
and depth. In the intertidal zone the density of this
species correlated positively with coarse sand,
carbonate and gravel contents, and negatively with
organic matter and fine sand contents (Table 2).
Among species relationships
In the subtidal zone, extensive overlap was observed
among the distribution of /iatella arctica. Venus
verrucosa and Fenerupis senegalensis (Fig. SA), and
similarly among the distribution of Abra alba, Thvasira
Bivalves and relatronships with environmental parameters
OLABARRIA & al.
flexuosa and Mvsella bidentata (Fig. 5B). though note
that the latter was very common throughout the
subtidal zone. Zhyvasira flexuosa and Dosinia exoleta
were not found together (Fig. 5B), since D. exoleta
occurs on infralittoral gravel and 7: flexuosa on muddy
sands or muds. Papillicardium papillosum showed high
densities (up to 150 individuals/m?) on the decpest
bottoms dominated by coarse sands.
In the intertidal Zone, extensive overlap was
observed between Fiatella arctica and Venus
verrucosa and between Parvicardium exiguum and
l'enerupis senegalensis (Fig. 5A). V. verrucosa and FH.
arctica occurred in the same types of habitat (coarse
sands and gravels), whereas ?. exiguum and
senegalensis both occurred on a wide range of bottom
types. C'erastoderma edule occurred most commonly at
intermediate tidal levels, with the highest densities at
SUBTIDAL
.385 .35 2: 2 5 53
Le STE 3 < 3
Species
Mytilus edulis 0.048
0.025
Thyasira flexuosa 0.584**
Mysella bidentata k
Papillicardium papillosum S 0.443**
Parvicardium exiguum -0.048°"
Cerastoderma edule
Abra alba
05262 ES ;
Venus verrucosa
!
=
©
a
à
Dosinia exoleta
INTERTIDAL
MS FS oM CA Si D/H
: ; -0.068
0.104" 0.048
Venerupis senegalensis
Te c
Hiatella arctica 0.471**
Species
Mytilus edulis
Thyasira flexuosa (HEERE
0.060 TS -0.357*
0.383*
002 RIFO0OS PE) ROTOE
0.250 -0.369 *
01522
Mysella bidentata 0.233 -
Papillicardium papillosum
Parvicardium exiguum
Cerastoderma edule
-0.020
0.086
0.052 0.000
+
0.201
0.008 TS -0.085
1 RS
-0.214 0.082 0.185 0:012%
Abra alba
Venus verrucosa
Dosinia exoleta
0.178 5 | 0161 5 | -0.145 5 | 025958
|
0.119 | 00377 | 0119 | 0,06
0.201
0.244 N° 0.011
0.518** | 02147 -0.397* -0.319* 0.130
Table 2. Coefficients of rank correlation (T-<) between densities of the different species (number of individuals in the
sample) and physicochemical factors (G : gravel content, CS : coarse sand content, MS : medium sand content,
FS: fine sand content, SC : silt-ciay content, OM : organic matter content, CA : carbonate content, S. : sorting
coefficient, D/H : depth/height with respect to tidal zero) NS : Not significant (p> 0,05), * p< 0,05, ** p< 0,01
V'enerupis senegalensis 020504001610 00000 21 os 0002287 -0.577**
[01922
Hiatella arctica
2c
6
OLABARRIA & al. Bivalves and relationships with environmental parameters APEX 13(3): 81-93, 20 juil. 1998
Fig. 3. Estimated densities (individuals per m2) of Parvicardium exiguum (A), Cerastoderma edule (B), Abra alba
(C) and Venus verrucosa (D) at sites at which that species was found.
87
APEX 13(3): 81-93, 20 juil. 1998 Bivalves and relationships with environmental parameters OLABARRIA & al
factor (possibly related to the positive correlation
observed between carbonate contents and particle size;
T, = 0.478, p < 0.01). Anyway A1. bidentata, C. edule
and !° senegalensis seemed to be less specific in their
choice of sediment types.
between 0.10 m and 0.94 m. Afvtilus edulis was found
in coarse sands and gravels, at densities of up to 100
individuals/m?
Generally speaking, the factors most strongly
influencing bivalve distribution were grain size and
depth. though carbonate content was also an important
Species Species
Nucula nitida Sowerby, 1833 Parvicardium nodosum (Turton, 1822)
Arca tetragona Poli, 1795 Cerastoderma edule (Linnaeus, 1758)
Striarca lactea (Linnaeus, 1758) Spisula elliptica (Brown, 1827)
Mhytilus edulis galloprovincialis Lamarck, 1818 Lutraria lutraria (Linnaeus, 1758)
Musculus subpictus (Cantraine, 1835) Phaxas pellucidus (Pennant, 1977)
Moerella donacina (Linnaeus, 1758)
Moerella pusilla (Phihippi, 1836)
Gobraeus tellinella (Lamarck, 1818)
Gobraeus depressa (Pennant, 1777)
Scrobicularia plana (da Costa, 1778)
Abra nitida (O.F.Müller, 1776)
Abra alba (Wood, 1802)
Venus verrucosa Linnaeus, 1758
Pecten maximus (Linnaeus, 1758)
Anomia ephippium Linnaeus, 1758
Pododesmus squamula (Linnaeus, 1758)
Monia patelliformis (Linnaeus, 1767)
Ostrea edulis Linnaeus, 1758
Crassostrea gigas (Thunberg, 1793)
Pisidium casertanum (Poli, 1791)
Loripes lacteus (Linnaeus, 1758)
Lucinoma borealis (Linnaeus, 1767) Gouldia minima (Montagu, 1803)
Myrtea spinifera (Montagu, 1803) Dosinia exoleta (Linnaeus, 1758)
Lucinella divaricata (Linnaeus, 1758) Tapes decussatus (Linnaeus, 1758)
Thyasira flexuosa (Montagu, 1803)
Lasaea rubra (Montagu, 1803)
Venerupis rhomboides (Pennant, 1777)
Venerupis saxatilis (Fleuriau de Bellevue, 1802)
Venerupis senegalensis (Gmelin, 1791)
Chamelea striatula (da Costa, 1778)
Clausinella fasciata (da Costa, 1778)
Timoclea ovata (Pennant, 1777)
Corbula gibba (Ohivi, 1792)
Hiatella arctica (Linnaeus, 1767)
Kellia suborbicularis (Montagu, 1803)
Montacuta substriata (Montagu, 1808)
Mpysella bidentata (Montagu, 1803)
Digitaria digitaria (Linnaeus, 1758)
Goodallia triangularis (Montagu, 1803)
Acanthocardia paucicostata (Sowerby, 1841)
Papillicardium papillosum (Pol, 1795) Nototeredo norvegica (Splenger, 1792)
Parvicardium exiguum (Gmelin, 1791) Thracia papyracea (Poli, 1791)
Table 3. List of the species detected in the present study showing overall relative abundance (i.e. percentage of the
total number of individuals found in all samples).
In our study area, the most important environmental
factors affecting the distribution of bivalves were
particle size and depth/ height. The current regime was
not investigated, though it 1s clearly determinant of the
DISCUSSION
Interpretation of spatial variations in the abundance of
benthic species 1s difficult, in view of the large number
of environmental factors which may act on benthic
communities. PEARSON & ROSENBERG (1978) studied
the factors involved in structuring the marine benthos,
and highlighted the importance of food availability as a
determinant of community structure. They concluded
that depth, latitude and water current speed are the
factors which have the strongest effect on food
availability. PETERSON (1979) reported that the factors
affecting abundance in the benthos may be divided into
density-dependent factors (such as competition.
predation, and adult-larva interactions) and the physical
properties of the sediment.
88
distribution of sediment types.
The dominant species in the intertidal area, Mytilus
edulis, Parvicardium exiguum and Cerastoderma
edule, showed a relationship with physicochemical
factors that was consistent with their autoecology. M.
edulis, which feeds on suspended detritus and
phytoplankton, is found primarily on coarse sediments
that allow it to attach itself by its byssus (TEBBLE,
1966). In the present study, the density of this species
correlated positively with coarse sediment content and
negatively with fine sand content. Both P. exiguum and
OLABARRIA & al. Bivalves and relationships with environmental parameters APEX 13(3): 81-93, 20 juil. 1998
Fig. 4. Estimated densities (individuals per m2) of Dosinia exoleta (A), Venerupis senegalensis (B), and Hiatella
arctica (C) at sites at which that species was found.
89
APEX 13(3): 81-93, 20 juil. 1998
C. edule are highly tolerant of variations in salinity,
and are typically indifferent to the type of substrate
(TEBBLE, 1966), the most important factors governing
their distnbution are the emersion time and water
current speed, both related to food availability
(FIGUERAS, 1956), since they are filter-feeders that can
only feed when submerged (LABORDA, 1986). In the
present study we found that the densities of both
species in the intertidal zone were negatively correlated
with height with respect to tidal level zero, while there
were no significant correlations with the other
environmental parameters. lenerupis senegalensis was
also abundant in the intertidal domain, with the most
important factor in its distribution being height: like
most venerids, it 1s adapted to the lower levels of the
interudal zone or to the subtidal zone. The importance
of tidal level and zoning has been discussed at length
by other authors (WOLFF, 1973; RAFAELLI & BOYLE.
1986; JUNOY & VIEITEZ, 1990). The distribution of this
species did not show any clear relationship with
sediment characteristics, in accordance with previous
reports: TEBBLE (1966), for example, found that tlus
species that inhabits all kinds of bottoms (sand, sandy
gravel, silty gravel, and silty sand), while in Kilkieran
Bay in Ireland, KEEGAN (1974) reported the presence
of this species in a great variety of biotopes. including
bottoms with clean sand, silty sand, maërl and on rare
occasions even conchiferous gravel. However, a study
by MORA (1980) in the Ria de Arousa (in southern
Galicia), found this species to be limited to clean sand
and gravel.
Of the donunant species in the subtidal zone. A/vsella
bidentata had a broad distribution throughout the inlet.
with high densities of individuals. The density of this
species in the subtidal zone showed no correlation with
any of the physicochemical parameters in accordance
with the fact that it is a highly ubiquitous species. able
to exploit a wide variety of environments, from sandy
to those with high contents of fine particles. It is also
characterized by its feeding behavior. which changes
depending on developmental stage: juveniles are
deposit-feeders while adults are filter-feeders
(OCKELMANN & MUUS, 1978). Thyasira flexuosa
showed a strong positive correlation with fine sand and
organic matter contents: similarly, other authors such
as LOPEZ-JAMAR et al. (1987) have reported that this
species 1nhabits silty sediments with à relatively high
organic matter content. This species has morphological
adaptations that prevent clogging of the branchial filter
by large particles in suspension (ALLEN. 1958).
Endosymbiont bacteria, which probably contribute to
its diet. are found in the gills (DANDO ef al.. 1985). In
our study area, the density of Abra alba correlated
positively with the sorting coefficient (which indicates
that it appears in poorly sorted sediments, with a wide
diversity of particles) and to a lesser extent with
organic matter content. GLEMAREC (1973) reports on
the presence of this species in heterogencous silty
facies. According to DAUVIN & GENTIL (1989), it is
plentiful in silt and sand sediments. tolcrates
physicochemical changes in the sediment very well.
90
Bivalves and relationships With environmental parameters
OLABARRIA & al.
and 1s rapidly adaptable (strategist r). LANDE (1975)
highlights the presence of this species in heterogeneous
sediments, and CORNET (1985) states that it can live on
highly varied bottoms. and that substrate granulometry
is not a decisive factor, although it requires a layer of
suspended detritus in the water-sediment interphase as
a source of food (GLEMAREC, 1964).
Venus verrucosa, Dosinia exoleta and Hiatella
arctica are species that live on bottoms characterized
by coarse sand, silt gravel or conchiferous gravel
(TEBBLE, 1966). In our study area, they occurred
largely on coarse sand and gravel, and the densities of
these species were negatively correlated with fine sand
content.
The observed correlations between the densities of
the different species and particle size and organic
matter content appear to be related to feeding
behaviour. For example, the densities of 7. flexuosa
and À. alba, which are burrowing detritus feeders,
correlated positively with organic matter and fine sand
contents. whereas the remaining species (with the
exception of Af bidentata which may change its
feeding mechanism depending on its developmental
stage) fed on particles in suspension, so that their
distribution correlates positively with sediments having
a larger particle size. According to CORNET (1986).
filter-fecders take over from detritus-feeders species
when the proportion of fine particles decreases.
LE VINTON (1977) reports that deposit-feeders dominate
in ecosystems with fine, soft sediments and that their
presence 1s linked to the silt-clay fraction, although this
latter has not been verified in our study area. By
contrast. filter-feeders dominate in sandy sediments
(SANDERS, 1958) and their distribution may be
governed more by hydrodynamic processes, which
determine sediment characteristics. than by the
characteristics of the sediment itself.
The marked overlap in the distribution of À. alba and
T. flexuosa is as expected given that these species
characteristically form part of a well-defined subtidal
zone community (see THORSON, 1957). Afysella
bidentata. which likewise showed considerable overlap
with these two species, 1s not characteristic of this
community but 1s ubiquitous and broadly distributed in
our study area. iatella arctica, V! verrucosa and F.
senegalensis, similarly showed overlapping
distributions, and all three occurred at high densities on
a maërl bed near Punta Redonda. This bed is made up
of ZLithothamniom corallioides and Phymatolithon
calcareum, on a shell-gravel bottom with a small
amount of silt. According to URGORRI et al. (1992).
these bottoms offer stable substrates which provide
good shelter for many species of small molluscs
including the juveniles of certain species. so that the
maërl acts as a hatchery.
The similar distributions in some areas of ?. exiguum
and l° senegalensis may be explained by the fact that
these species are characteristic of the intertidal facies
occupicd by Zostera noltii within the limited
community Of A/acoma balthica (THORSON., 1957).
APEX 13(3): 81-93, 20 juil. 1998
Bivalves and relationships with environmental parameters
OLABARRIA & al.
O Hiatella arctica E Venerupis senegalensis E Venus verrucosa M Parvicardium exiguum
CCD]
nn
CN NN]
sal2ods %
Intertidal stations
Subtidal stations
O Mysella bidentata
E Abra alba
O Dosinia exoleta
M 7hyasira flexuosa
222 NN
TP PPT PTE TEE TT
CPP NN)
TT
sa12ads %
Intertidal stations
Subtidal stations
Venerupis
Venus verrucosa,
Û
Fig. 5. Plots illustrating the overlaps in the distributions of Hiatella arctica
Senegalensis and Parvicardium exiguum (A) and Abra alba, Mysella bidentata, Thyasira flexuosa and Dosinia
exoleta (B). The horizontal axis shows sample number. The vertical axis shows relative abundance, here defined as
the number of individuals in that sample expressed as a percentage of the maximum number of individuals
sample recorded for that species
per
91
APEX 13(3} 81-93, 20 juil 1998
This has likewise been reported by CURRAS & MORA
(1991) in the Ria de Ribadeo (likewise on the north
coast of northwest Spain), where P. exiguum and F
senegalensis were found on muddy sand or sandy mud
bottoms covered by Z. noltii. This phanerogam gives
nise 10 a more diversified habitat, and its rhizomes and
roots compact the sediment and provide protection
from predators (ECKMAN, 1987). Both population
densities and species richness are thus typically higher.
Finally, /7. arctica and F. verrucosa showed closely
overlapping distnbutions since both occurred in the
dumping area of the dredging operations carried out in
the channel of the ria. The dumped material 1s
characterized by coarse gravel and conchiferous gravel,
which are typical habitats for these species.
ACKNOWLEDGEMENTS. This research form part of
project XUGA 20005B95, and was partially financed
by a Predoctoral Grant from the Xunta de Galicia.
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25
HOUART
Eight new species of Muricidae
Description of eight new species of Muricidae (Gastropoda)
Roland HOUART
Research associate, Institut roval des Sciences naturelles de Belgique
Département des Invertébrés Récents
Rue Vautier, 29, B-1000 Brussels, Belgium
KEY WORDS. Gastropoda, Muricidae, new species.
ABSTRACT. The following new species of Muricidae are described, and compared with related
species: Aftiliosa edingeri and Favartia eastorum from Western Australia, Favartia deynzeri from the
Red Sea, Apixystus rippingalei from Queensland, 7rophonopsis bassetti from New South Wales and
Queensland, Orania rosadoi from Mozambique, Ergalatax dattilioi from the Philippine Islands,
Indonesia, and Japan, and Thaïs herberti from South Africa.
RESUME. Les espèces suivantes sont décrites et comparées avec des espèces apparentées: Aftiliosa
edingeri et Favartia eastorum d'Australie Occidentale, Favartia deynzeri de la Mer Rouge, Apixystus
rippingalei du Queensland, 7rophonopsis bassetti de la Nouvelle-Galles du Sud et du Queensland,
Orania rosadoi de Mozambique, Ergalatax dattilioi des Philippines, d'Indonésie et du Japon, et Zhais
APEXx 13(3): 95-109, 20 juil. 1998
herberti d'Afrique du Sud.
INTRODUCTION
New species of Muricidae continue to be discovered
thanks to dredging programs, expeditions, and private
collecting. The eight new species described here
originate from different sources: Attiliosa edingeri,
Favartia eastorum, F. deynzeri and Orania rosadoi
were recently discovered by private collectors:
Apixystus rippingalei and Trophonopsis bassetti were
collected during several cruises by HMAS Kimbla
(1976, 1977, 1984), and FRV Kapala (1975, 1977,
1979) (material housed in Australian Museum.
Sydney), Ergalatax dattilioi by private collectors, and
during the MUSORSTOM 1! expedition (1976), by
R.V. Vauban off the Philippine Islands, and the
KARUBAR expedition (1991), by R.V. Baruna Java 1
in the Banda and Arafura Seas, and off the Kai and
Tanimbar Islands, Eastern Indonesia (material housed
in Muséum National d'Histoire Naturelle, Paris): Thais
herberti during the Natal Museum Dredging Program,
1989 (Natal Museum, Pietermaritzburg).
Abbreviations
AMS: The Australian Museum, Sydney. Australia.
BMNH: The Natural History Museum. London. Great
Britain.
IRSNB:
Belgique.
MNAN: Muséum National d'Histoire Naturelle. Paris.
France.
NM: Natal Museum, Pietermaritzburg. South Africa.
Institut royal des Sciences naturelles de
NMNZ: Museum of New Zealand, Wellington.
QM: Queensland Museum, Brisbane, Australia.
WAM: Western Australian Museum, Perth, Australia.
dd: empty shell.
Iv.: live-taken specimen.
SYSTEMATIC ACCOUNT
Family MURICIDAE Rafinesque, 1815
Subfamily MURICINAE Rafinesque, 1815
Genus Afttiliosa Emerson, 1968
Type species (by original designation): Coralliophila
incompta Berry, 1960 (= Peristernia nodulosa A.
Adams. 1854); Recent. Eastern Pacific.
Remarks
When I described Aftiliosa goreensis (HOUART, 1993)
from Senegal, I recorded that four species and one
subspecies of Attiliosa were known at that time: À.
nodulosa (A. Adams. 1855) from the Eastern Pacific:
A. nodulifera (Sowerby. 1841), and its subspecies
caledonica Jousseaume, 1881 from the Indo-West
Pacific; À. philippiana (Dall, 1889) from Florida, and
A. aldridgei. known from different localities in the
Western Atlantic. Since then, in addition to À.
goreensis. Other species of Artiliosa have been
95
APEX 13(3} 95-109, 20 ul 1998
Eight new species of Municidae
HOUART
described: À. bozzettii Houart, 1993 from Somalia, À
glenduffvi Petuch, 1993 from the Dominican Republic,
and À. ruthae Houart, 1996 from the Philippine Islands
Moreover, in 1993 I neglected to mention À. eorri
(Cernohorsky, 1976) from the Andaman Islands
Since the description of À. goreensis, I have had the
opportunity to examine additional specimens of À
nodulifera caledonica, including a specimen from
Mindanao, Philippine Islands (Fig. 31). The shell 1s
large, almost 28 mm in length and dark brown while
other specimens of A. caledonica are usually white.
However, apart from colour, and size, I cannot separate
it from typical specimens of À. caledonica. Since À.
nodulifera lives throughout the Philippine Islands, 1t
seems now that À caledonica should be raised to
specific rank. However, À. nodulifera 1s not yet
recorded from other localities where À. caledonica
occurs, such as New Caledonia and Polynesia
(TRÔONDLE & HOUART, 1992).
Attiliosa edingeri n sp.
Figs 1-4, 40
Type material examined.
Western Australia: off Esperance, 34-36 m. holotype
WAM S.IIOI and 1 paratype WAM S.1102: off
Esperance, amongst weed, sand and rubble, 31 m., 1
paratype coll. A. Edinger, 1 paratype coll. R. Houart
(all 1v.).
Distribution.
Off Esperance, Western Australia, taken alive at 31-36
m.
Description.
Shell biconical, large for the genus, up to 35.7 mm in
length at maturity; hightly built, squamous. Spire high.
with 1.5 protoconch whorls and up to 6 broad, convex,
weakly shouldered teleoconch whorls. Suture
adpressed. Protoconch small, whorls rounded, smooth:
terminal varix unknown (eroded). Axial sculpture of
teleoconch whorls consisting of low, rounded.
squamous ribs and of numerous growth lamellae. First
whorl with 11 ribs, second to fifth with 11 or 12 ribs.
last whorl with 9 nbs. Spiral sculpture of weak,
squamous, rounded, primary and secondary cords and
some weak, squamous threads. First whorl with 2 cords
and 2 or 3 threads on shoulder., second to fourth with 2
or 3 cords and 3 or 4 threads on shoulder, penultimate
with 3 or 4 cords and 3-6 threads on shoulder, last adult
whorl with 12-16 cords and 44 shallow, squamous
threads on shoulder.
Aperture large, ovate; columellar lip smooth; lip
weakly erect partially, adherent at adapical extremity;
anal notch shallow, broad; outer lip crenulate, with 7
weak, elongate denticles within, denticles occasionally
absent. Siphonal canal short, broad, strongly
96
abaperturally recurved at extremity, with 2 or 3
squamous spiral cords and single thread.
Light tan or orange with darker coloured spiral band
on periphery of last whorl, occasionally lighter
coloured axial ribs. Aperture white.
Operculum dark brown, ovate, with terminal nucleus.
Radula with a long, broad central cusp: narrow, long,
lateral cusps with broad base. Lateral teeth sickle
shaped, broad.
Remarks.
Attiliosa edingeri n.sp. is highly distinctive from any
known Indo-Pacific species of Artiliosa. It differs from
A. nodulifera, À. caledonica, À. ruthae and À. orri in
having à spineless shell, and from these species and À.
bozzettii, in having more conspicuous, numerous, and
squamous spiral cords and threads, a smooth
columellar lip, and à broader aperture relative to the
shell length. It is currently the largest known Recent
species of Artiliosa.
Etymology.
Named for Andrew Edinger, Mandurah, Western
Australia, who, together with Peter Clarckson,
discovered the habitat of the new species, and sent it to
me for identification.
Subfamily MURICOPSINAE
Radwin & D'Attilio, 1971
Genus Favartia Jousseaume, 1880
Type species (by original designation): AMurex
breviculus Sowerby, 1834: Recent, Indo-West Pacific.
Favartia eastorum n sp.
Figs 5-7
Type material examined.
Western Australia: Peak Island, N of Exmouth Gulf,
WA: 21°36'S, 114°36'E, 24-27 m, buried in silty sand
pockets, shelving limestone reef with sparse covering
of small sponges, gorgonians, and hard corals, holotype
WAM S.1103; paratypes 1 WAM S.1104: 1 AMS C.
203326; 1 coll. R. Houart (all Iv.).
Distribution.
Peak Island, N of Exmouth, Western Australia, taken
alive at 24-27 m.
Description.
Shell of medium size for the genus, up to 21.5 mm in
length at maturity, tuberculate. Spire high. up to 5
weakly convex, shouldered teleoconch whorls, suture
Eight new species of Muricidae APEX 13(3): 95-109, 20 juil. 1998
Figs 1-4. Attiliosa edingeri n.sp. Figs 1-2. Western Australia, off Esperance, 34-36 m, holotype WAM S.1101, 31.9
mm. Fig. 3. Western Australia, off Esperance, 34-36 m, paratype WAM S.1102, 30.3 mm. Fig. 4. Western Australia,
off Esperance, 31 m, paratype coll. Edinger, 35.7 mm. Figs 5-6. Favartia eastorum n.sp., Western Australia, Peak
Island, N of Exmouth Gulf, WA, 21°36'S, 114°36'E, 24-27 m, holotype WAM S.1103, 21.4 mm.
97
APEX 13(3) 95-109, 20 juil. 1998
adpressed. Protoconch unknown (eroded). Axial
sculpture of teleoconch whorls consisung of high,
strong, broad, rounded varices: first and second whorls
with $ or 6 varices, third and fourth with 4 or 5, last
whorl with 4 vances. Spiral sculpture of high, strong,
nodose primary cords, secondary cords on shoulder and
siphonal canal: spiral sculpture of first whorl eroded,
second with 2 cords on whorl, 2 on shoulder, third with
one on whorl, 2 or 3 on shoulder, fourth with 2 on
whorl, 2 on shoulder, last whorl with 5 on whorl and 3
on shoulder, cords more strongly developed on axial
varices, shallow or/and eroded on early whorls.
Aperture small, ovate, columellar lip flanng, narrow,
smooth, nm partially erect, adherent at adapical
extremity, anal notch shallow, broad: outer lip erect,
crenulate, smooth within. Siphonal canal short, broad.
abaperturally recurved at extremity, narrowly open,
with 2 or 3 smooth, or almost obsolete spiral cords.
Light tan with darker coloured blotches, mostly on
spiral cords. Aperture white.
Operculum dark brown, ovate-elongate with terminal
nucleus. Radula unknown.
Remarks.
Favartia confusa Brazier, 1877 (Figs 8-9) differs in
having a larger shell relative to the number of
teleoconch whorls. Furthermore, in Æ confusa the
siphonal canal is markedly straighter, the spiral cords
are narrower, the varices are broader and flanged
adapically and abapically; abapical flange almost
smooth, extending up to the extremity of the siphonal
canal.
Favartia cyclostoma (Sowerby, 1841) has a broader,
more shouldered shell with more numerous, narrower
spiral cords, more fimbriated varices, and a more
rounded aperture.
Etymology.
At the request of Peter Clarckson. the species 1s named
for Milton and Aileen East, of Geraldton, West
Australia, who first showed him a specimen from their
collection.
Favartia deynzeri n sp.
Figs 10-13
Type material examined.
Red Sea: Egypt. Shab Shareer, 15-20 m. holotype
IRSNB IG 28515/478. Paratypes: 1 coll. A. Deynzer; 1
coll. R. Houart (all Iv.).
Distribution.
Shab Shareer, Egypt, taken alive at 15-20 m.
98
Fight new species of Muricidae
HOUAR1T
Description.
Shell medium sized for the genus, up to 17.18 mm in
length at maturity (holotype), heavy, moderately
spinose. Spire high with 1.5 protoconch whorls and up
to 5 broad, angulate, weakly shouldered teleoconch
whorls. Suture impressed, partially obscured by small
axial lamellae. Protoconch small, giobose, whorls
rounded: terminal varix unknown (eroded). Axial
sculpture of teleoconch whorls consisting of 4 or 5
high, strong, broad, rounded varices from first to last
whorl; varices more developed on shoulder. Spiral
sculpture of high, strong, squamous cords: two spiral
cords on early whorls; last whorl with 5 cords, ending
as short spinelike projections on varices, last whorl
occasionally with minor spiral threads.
Aperture small, ovate;, columellar lip narrow, flaring,
smooth, rm partially erect, adherent at adapical
extremutv, anal notch obsolete, outer lip erect,
crenulate, with 4 or 5 weak, occasionally obsolete, lirac
within. Siphonal canal moderately long, broad, straight.
abaperturally recurved at extremity, narrowly open,
with 3 or 4 frondose, short spinelets.
Creamy-white, occasionally with light tan blotches
on spiral cords or shoulder; aperture white.
Operculum dark brown, ovate, with terminal nucleus.
Radula unknown.
Remarks.
Favartia deynzeri is highly distinctive from any other
Indo-West Pacific or Red Sea species of Favartia by its
form, sculpture and particular ornamentation of the
siphonal canal. F: cyclostoma (Sowerby, 1841) and F
sykesi (Preston, 1904) which superficially resemble F
deynzeri, have a larger and broader shell, with more
numerous, narrower varices, and narrower, larger
siphonal canal with different ornamentation.
Etymology.
Named for AI Deynzer, Sanibel, Florida, who sent me
the shells for identification.
Subfamily TROPHONINAE (sensu lato)
Cossmann, 1903
Genus Apixystus Iredale, 1929
Type species (by original designation): 7rophon
stimuleus Hedley, 1907; Recent, North-East Australia.
Apixystus rippingalei n sp.
Figs 14-16, 46
Type material examined.
Queensland: E. of Lady Musgrave Island, 23°52.5' -
HOUART Eight new species of Muricidae APExX 13(3): 95-109, 20 juil. 1998
Fig. 7. Favartia eastorum n.sp. Western Australia, Peak Island, N of Exmouth Gulf, WA, 21°36'S, 114°36'E, 24-27
m, paratype coll. R. Houart, 18.8 mm. Figs 8-9. Favartia confusa (Brazier, 1877). Australia, Queensland, Darnley
Id, Torres Strait, 55 m, holotype AMS C.077183, 26.8 mm. Figs 10-13. Favartia deynzeri n.sp. Red Sea, Egypt,
Shab Shareer, 15-20 m. Figs 10-11. Holotype IRSNB 1G 28515/478, 17.2 mm. Fig. 12. Paratype coll. R. Houart, 13
mm. Fig. 13. Paratype coll. A. Deynzer, 15.6 mm.
99
APEX 13(3) 95-109, 20 juil. 1998
Eight new species of Muricidae
HOUARI
23°51.9' S, 152°42.7' - 152°41.7' E. 296 m. holotype
AMS C.313232 (dd)
Paratypes. 42 AMS C.313230. 1 BMNH 1996286. I
MNAHN. 1 NM L4346/T1519, 1 NMNZ M272617. |
QM MO 61758. 2 coll. R. Houart (all dd)
Other material examined.
Queensland: SE of Swain Recfs. 22°26.27' - 22°202'S,
153°17.13"- 152°176 E 187 m AMS C 321907 (7):
Capricom Channel, 23°8.6' S, 152°16.6' E, 155 m.
AMS C.321954 (1): E of North West Is. Capricorn
Channel, 23°15.2' S, 152°24.l' E, 284 m, AMS
C.321963 (1), 245 ml E of Lady Musgrave Island.
23°33.7'S, 152°37'E. 339-348 m. AMS C.125294 (3):
of Sandy Cape, 24°43.5' - 24°438' S, 153°33.4 -
153°333'2E, 604: :m, “AMS C313229-0 (1); rot
Maroochydore, 26°41.2' S. 153°38.4'E, 200 m, AMS
C.321943, (1), off Cape Moreton, 27°0' S, 153°34' -
153936" E, 128-183 m. AMS C.150076., (3) (all dd).
Distribution.
Queensland, Australia, in 128-604 m (Fig. 46).
Description.
Shell medium sized for the genus, up to 44 mm in
length, spinose, delicate. Spire high with 1.5-1.75
protoconch whorls and up to 4 angulate, shouldered,
spinose, telcoconch whorls. Suture impressed.
Protoconch large, globose, whorls rounded, smooth:
terminal varix shallow, delicate, thin, weakly curved.
Axial sculpture of teleoconch whorls consisting of
narTOW, spinose lamellae: first whorl with 7 or 8
lamellae, second with 10 or 11, third with 12-14, last
whorl with 14 lamellae. Spiral sculpture of low, weak,
smooth cords on first teleoconch whorl, and of high.
rounded cords on other whorls, ending as short spines;
shoulder spines more conspicuous. First and second
whorl with 2 or 3 cords, third with 3 or 4 cords, last
whorl with 4 cords.
Aperture moderately small, rounded: columellar lip
flanng, smooth, rim partially erect, adherent at adapical
extremity; anal notch obsolete; outer lip undulate, with
4 weak or strong, elongate denticles within. Siphonal
canal moderately long. 16-23 % of total shell length:
open, smooth or occasionally with a single, smooth.
medial spiral cord.
Transluscent white.
Radula and operculum unknown.
Remarks.
The shell sculpture in specics of .Ipixystus 1s
interspecifically and intraspecifically rather uniform.
Apixvstus stimuleus (Hcdley, 1907) (Fig. 17) has 9 or
10, less frilly axial lamellac on the last telcoconch
whorl, the shoulder spines are broader and shorter;
there are 2 spiral cords on the penultimate whorl
instcad of 3, and 2 or 3 low spiral cords on the last
whorl. instead of the 4 strong, high cords. À. leptos
Houart, 1995 (Figs 18-19) has 2 spiral cords on the
penultimate whorl, 2 on the last whorl, occasionally
with 2 low. shallow threads abapically: the spines are
shorter, and the siphonal canal is shorter and more
wcakly recurved. À. recurvatus (Verco. 1909) (Figs 20-
21) is relatively larger, less shouldered, with fewer,
lower axial lamellae, shorter spines, and lower spiral
sculpture.
Etymology.
Named in memory of the late Oswald (Ossie) H.
Rippingale, Margate Beach, Queensland, friend, artist
and shell lover.
Genus Trophonopsis
Bucquoy & Dautzenberg, 1882
Type species
muricatus Montagu.
North-East Atlantic.
(by original
1803,
designation): Afurex
Recent, Mediterranean.
Trophonopsis bassetti n.sp.
Figs 22-23, 47
Type material.
New South Wales: SE of Clarence River, 29°4l' -
29932" $S, 153°45 - 153°47' E, 405-412 m, holotype
AMS C.313223 (Iv.), and 1 paratype coll. R. Houart
(dd).
Other paraiypes: Queensland: E of Lady Musgrave
Island, 23°52.5' - 23°519°$S 152492 752 RIRE
296 m, 1 AMS C.313231 (Iv.); New South Wales: E of
Cape Three Points, 33°28' - 33°29'S, 152°4'- 15293'E,
457-476 m, 2 AMS C.321596 (dd); off Sydney, 34°4.2'
S, 151°37.4' E, 393 m, 1 AMS C.322783 (Iv.): off
Ulladulla, 35°30' - 35°33' $S, 150°48' - 150°47'E, 549
m, 1 AMS C.313224 (dd).
Figs 14-16. Apixystus rippingalei n.sp. Australia, Queensland, off Lady Musgrave Id, 296 m, Figs14-15. Holotype
AMS C.313232, 4 mm. Fig. 16. Paratype AMS C.313230, 44 mm. Fig. 17. A. stimuleus (Hedley, 1907). NSW,
Sydney, 22 miles east of Narrabeen, 146 m, holotype AMS C.25797, 3.3 mm. Figs 18-19. À /eptos Houart, 1995.
New Caledonia, holotype MNHN, 4.8 mm. Figs 20-21. À. recurvatus (Verco, 1909). NSW, off Sydney, AMS
C.150080, 7 mm. Figs 22-23. Trophonopsis bassetti n.sp. Australia, NSW, SE of Clarence River, 405-412 m,
holotype AMS C.313223, 89 mm. Fig. 24. 7Jrophonopsis plicilaminatus (Verco, 1909). South Australia, off
Beachport, 200 fms (365 m), paratype AMS C.31093, 15.9 mm. Fig. 25. Trophonopsis segmentatus (Verco, 1909).
South Australia, off Cape Jaffa, 90 fms (165 m), paratype AMS C.31065, 9.5 mm. Fig. 26. 7rophonopsis simplex
(Hedley, 1903). NSW, Sydney, off Port Hacking, coll. R. Houart, 11 mm.
100
HOUART Eight new species of Muricidae APEX 13(3): 95-109, 20 juil. 1998
101
APEX 13(3): 95-109. 20 juil 1998
Distribution.
New South Wales and Queensland, Australia, 296-549
m, taken alive in 296-393 m (Fig. 47)
Description.
Shell medium sized for the genus, up to 13.9 mm in
length at maturity (paratype AMS C.321596). slender,
weakly spinose, delicate. Spire High with 1.5
protoconch whorls and up to 4.5 shouldered, spinose
teleoconch whorls. Suture impressed. Protoconch large,
globose: whorls rounded, glossy; terminal vanix thin,
low, weakly curved. Axial sculpture of teleoconch
whorls consisting of low, weak lamellae. Lamellae
more strongly developed on shoulder, occasionally
producing short, spinelike projections: 7 or 8 lamellae
on first whorl, 10-12 on second, 11-13 on thurd. 11-15
on last whorl. Spiral sculpture of broad, rounded cords:
first, second and third whorl with 2 cords, last whorl
with 5: shoulder smooth, except axial lamellae,
occasionally with single, low. shallow cord on last
whorl.
Aperture moderately small, angulate: inner lip almost
horizontal, forming an angle of approximately 85-86°
with axis of shell: columellar lip smooth, adherent; anal
notch shallow; outer lip smooth, with 3 weak., low
denticles within; adapical denticle more apparent.
Siphonal canal medium sized or long, 29-33% of total
shell length, narrow, straight, or weakly abaxially
recurved. with 2 or 3, squamous, narrow spiral cords
adaperturally.
Uniformly milky-white.
Operculum and radula unknown.
Remarks.
Trophonopsis segmentatus (Verco, 1909) (Fig. 25) has
a more convex shell with rounded, crowded spiral
cords, more numerous cords on spire whorls, and a
smoother shoulder. The spire is higher, and the
siphonal canal is markedly shorter, occupying 18% of
the total shell length in 7: segmentatus instead of 29-
33%. The aperture of 7. segmentatus is ovate, with a
broad columellar lip, instead of triangular with a
narrow columellar lip.
Trophonopsis plicilaminatus (Verco, 1909) (Fig. 24)
has fewer, lower spiral cords, a smooth shoulder, and
an ovate aperture.
Another species, described from New South Wales,
Trophonopsis simplex (Hedley, 1903) (Fig. 26) has à
smoother, more convex shell with a roundly-ovate
aperture. The spire whorls are rounded with more
numerous spiral cords (8 or 10 cords on last whorl with
a higher density on abapical part of the whorl).
Etymology.
Named in memory of the late Arthur Bassett, Shark
Bay. West Australia, whom I have known for many
102
Eight new species of Muricidae
HOUART
years, and who has sent me many Australian muricids
and other Australian shells
Subfamily ERGALATAXINAE
Kuroda & Habe, 1971
Genus Orania Pallary, 1900
Type species (by original designation): Pseudomurex
spadae Libassi, 1889 (= Murex fusulus Brocchi, 1814);
Pliocene to Recent, Mediterranean, eastern Atlantic.
Orania rosadoi n sp.
Figs 27-29, 43
Type material examined.
South Mozambique: Quissico area, ex pisce, in 50-60
m, holotype NM L3374/T1463. Paratypes: 6 NM
L3581/T1464: 1 MNHN: 2 coll. J. Rosado of Maputo,
1 coll R. Houart (all dd).
Distribution.
Quissico area. South Mozambique. 50-60 m (ex pisce).
Description.
Shell medium sized for the genus, up to 13.3 mm in
length at maturity (paratype NM). slender, weakly
spinose, lightly built. Spire high with 1.5 protoconch
whorls and up to 5 angulate, shouldered, weakly
spinose teleoconch whorls. Suture impressed.
Protoconch small, globose; whorls rounded, smooth;
terminal varix shallow, thin, weakly curved. Axial
sculpture of teleoconch whorls consisting of low.
broad, nodose, weakly spinose varices: 8-10 varices
from first to penultimate whorl; 7 or 8 on last whorl.
Spiral sculpture of high, squamous, primary and
secondary cords: 2 on first whorl, 2 or 3 on second and
third, 3 or 4 on fourth, 4 or 5 primary cords and 2-4
secondary cords on last whorl; carinal cord forming
small, open spines at intersection with axial varices.
Aperture moderately large, narrow, ovate: columellar
lip smooth, rim broken in all specimens: anal notch
shallow, broad; outer lip with 6 or 7 elongate denticles
within. Siphonal canal short. straight. open, with 3 or 4
spiral cords.
Light brown with darker coloured blotches on suture,
and occasionally with darker coloured secondary spiral
cords between third and fourth abapical primary cords
of last whorl.
Operculum and radula unknown.
Remarks.
The classification of this new species in Ergalataxinae,
and in Orania, 1s tentative, as no radula is available.
HOUART Eight new species of Muricidae APEX 13(3): 95-109, 20 juil. 1998
Figs 27-29. Orania rosadoi n.sp. Mozambique, Quissico area. Figs 27-28. Holotype NM L3374/11463, 11.9 mm.
Fig. 29. Paratype NM L3581/171464, 10.5 mm. Fig. 30. Ergalatax dattilioi n.sp. Indonesia, Tanimbar Ids, 184-186 m,
MNHN, 14.1 mm. Fig. 31. Aftiliosa caledonica (Jousseaume, 1881). Philippine Islands, Mindanao, Balut Id, 200 m,
coll. A. Deynzer, 27.7 mm.
103
APEX 13(3): 95-109, 20 juil. 1998
Nevertheless, the size, shape, and sculpture of the shell
are similar to other species included in the
Ergalataxinae, and more particularly in Orania.
Another possibility might be laughtia Houart, 1996
(Ocenebrinae), currently known exclusively from
southern Africa. However, the shells of Faughtia
species are usually more rounded. with lower axial
sculpture or none, and a broader aperture.
Orania rosadoi n.sp. resembles ©. mixta Houart,
1995, but has fewer and broader, secondary spiral cords
on last teleoconch whorl, and à paucispiral protoconch
of 1.5 whorls, whereas ©. mixta has a multispiral,
conical protoconch of sinusigera type, with 3.5 whorls.
Adults of ©. rosadoiï also seems to have à constantly
smaller size than ©. mixta.
Orania rosadoi n.sp. differs from ©. adiastolos
Houart, 1995, another species with paucispiral
protoconch, currently known from Zululand (South
Africa), and New Caledonia, in having a narrower,
more shouldered shell, with fewer secondary spiral
cords on last whorl, and elongate denticles within the
outer lip, rather than lirae.
Etymology.
Named for Mr. J. Rosado, Maputo, who donated the
type material.
Genus Ergalatax Iredale, 1931
Type species (by original designation): Ærgalatax
recurrens Iredale, 1931 (= Murex pauper Watson.
1883), Recent, Indonesia.
Ergalatax dattilioi n sp.
Figs 30, 35-37, 44, 45
Cytharomorula sp. - HOUART, 1986: 432, pl. 5, fig. 19
(holotype illustrated).
Ergalatax tokugawai - HOUART, 1997: 290 (not
Ergalatax tokugawai Kuroda & Habe, 1971).
Type material examined
Philippine Islands: 13°57' N, 120°16' E, 150-159 m,
holotype MNHN (Iv.); Cebu, Mactan Island, 73-110 m.
in coral, 1 paratype coll. R. Houart (Iv.).
Other material examined.
Philippine Islands: Mactan Island, Punta Engaño,
approximately 50 m, coll. F.J. Springsteen (2 Iv.).
Indonesia: Tanimbar Islands. 07°59' S, 133°02'E, 184-
186 m, MNEHN (2 1v. 1 dd), 08°00'S, 132°59'E, 206-
210 m, MNEN (4 dd).
Japan: Wakayama Pref, Minabe, 100-120 m, coll. R.
Houart (1 1v.).
104
Eight new species of Muricidae
HOUART
Distribution.
Tanimbar, Indonesia, the Philippine Islands, and South
of Japan. taken alive at 50-186 m (Fig. 45)
Description.
Shell small sized for the genus, up to 17.7 mm in
length at maturity, slender, tuberculate. Spire high with
3+ protoconch whorls and up to 6 weakly angulate,
shouldered teleoconch whorls. Suture impressed.
Protoconch conical, acute, smooth, glossy, of
sinusigera type. Axial sculpture of teleoconch whorls
consisting of high, strong, nodose ribs: 8 on first whorl,
8 or 9 on second, 8-10 on third, 9-11 on fourth and
fifth, 7-9 on last whorl. Spiral sculpture of low, nodose,
broad cords, covered by narrow, occasionally shallow
threads: 2 or 3 cords on first whorl, 2 or 3 cords on
second whorl, and 1 thread on shoulder, 2 or 3 on third
with 1 thread and 3 or 4 threads on shoulder, 2 or 3 on
fourth with 4 threads and 4-6 threads on shoulder, 3 on
fifth with 16-18 threads and 8-12 threads on shoulder.
6-9 cords on last whorl with numerous threads and 12-
15 threads on shoulder.
Aperture large, ovate: columellar lip smooth,
occasionally with 1 or 2 low, weak knobs abapically.
Rim adherent, weakly erect abapically; anal notch
narrow, deep: outer lip erect, smooth, with 5 or 6 weak
or strong elongate denticles within. Siphonal canal
short, broadly open, with some narrow threads.
Light tan or light brown, occasionally with some
weakly darker coloured blotches on axial ribs, and
occasional darker coloured threads between spiral
cords. Aperture white.
Operculum and radula unknown.
Remarks.
Ergalatax dattilioi Was recently identified as E.
tokugawai (HOUART, 1997), but from comparison on
type material (holotype, Figs 38-39) they are clearly
distinct species. Æ dattilioi has more elongate
teleoconch whorls, and different spiral sculpture. In Æ
dattilioi the spiral cords are broad and covered with
narrow threads, in Æ tokugawai, the spiral cords are
narrow, smooth, with 2 or 3 threads between each pair
of cords. The spiral threads are more numerous, and
narrower in £. dattilioi.
The genus Cytharomorula, once used for the species
(HOUART, 1986), resembles Ergalatax although having
shells with more adpressed suture, higher last
teleoconch whorl, and shorter siphonal canal relative to
the shell length.
Etymology.
Named in memory of the late Anthony D'Attilio, San
Diego, California, known throughout the world for his
numerous papers on Muricoidea.
Eight new species of Muricidae APEX 13(3): 95-109, 20 juil. 1998
Figs 32-34. Thais (Mancinella) herberti n.sp. Figs 32-33. South Africa, northern Natal, off Glenton Reef, 110 m,
holotype NM S2603/T1465, 31.1 mm. Fig. 34. Protoconch and first whorls. Figs 35-36. Ergalatax dattilioi n.sp
Philippine Islands, 13°57' N, 120°16'E, 150-159 m, holotype MNHN, 16.2 mm
105
APEX 13(3): 95-109, 20 juil. 1998
Subfamily RAPANINAE Gray, 1853
Genus Thais Rôding, 1798
Subgenus Mancinella Link, 1807
Type species (by absolute tautonomy, ICZN, opinion
911): Mancinella aculeata Link, 1807 (= lolema
alouina Rôding, 1798); Recent, Indo-West Pacific.
Thais (Mancinella) herberti n.sp.
Figs 32-34, 41-42
Mancinella cf. siro (Kuroda, 1931) - LUSSI & BRINK.
1996: 3, fig. 17.
Type material examined.
South Africa: Northern Natal. off Glenton Reed.
29°13.7'"S, 32°020'E, 110 m, sponge. holotype NM
S2603/T1465 (lv.);, Natal, off Scottburgh, 100 m, 1
paratype NM B3485/T1466 (Iv.).
Other paratypes: off Port Shepstone, 70 m, eroded shell
+ sponge rubble, 1 NM B3665/T1467 (dd), Zululand,
NE. of Gipsy Hill, 27°45.2" S, 32°39.8' E, 110 m,
sponge, stones, 1 NM E3731/T1468 (Iv.): Zululand., off
Richards Bay, 28°59' $S, 32°11' E, 100 m, 1 NM
B6313/T1469 (lv.): 1 coll. R. Houart (lv.): Zululand,
off Matigulu R. mouth, 29°17.1'S, 31°503'E, 50 m,
mud, 1 NM VS5136/T1471 (Iv.); Zululand, off Matigulu
Bluff, 29°21.4' S, 31°562' E, 90 m. sponge. 1 NM
E8760/T1470 (Iv.).
Distribution.
Natal and Zululand, South Africa, taken alive at 50-110
m, on sponge bottoms on mid-continental shelf.
Description.
Shell small for the subgenus, up to 34 mm in length at
maturity (paratype NM E8760/T1470), heavy,
biconical. Spire high, with 2+ protoconch whorls
(partially broken), and up to 5 or 6 broad, convex,
shouldered teleoconch whorls. Suture adpressed.
Protoconch large, conical; whorls rounded, minutely
punctate, with a narrow keel abapically. Terminal varix
of sinusigera type. Axial sculpture of teleoconch
whorls consisting of low, broad, weakly spinose ribs,
each with one short, acute, broad spine on shoulder.
Last whorl with 2 rows of short spines. First whorl with
10-12 axial ribs, second with 10, third, fourth and fifth
with 9 or 10, last whorl with 7 or 8 ribs. Spiral
sculpture of high, strong, squamous, primary and
secondary cords: first whorl with 3 cords on whorl and
2 on shoulder, second with 3 on whorl, 4-6 on
shoulder, third and fourth with 6 on whorl, 9 on
shoulder, fifth whorl with 8 or 9 on whorl, 12 or 13 on
shoulder, last whorl with 27 on whorl, 14-17 on
shoulder.
106
Eight new species of Muricidae
HOUART
Aperture large, broad, ovate, columellar lip smooth,
adherent, weakly detached abapically; anal notch
broad; outer lip crenulate, with 8-10 strong lirae within.
Siphonal canal short, broad, straight, broadly open,
with $ or 6 primary and secondary, rounded spiral
cords.
Light tan, aperture white.
Operculum dark brown, D-shaped, with lateral
nucleus in center right. Radula with a long, broad
central cusp; lateral cusps with inner lateral denticles; 3
or 4 marginal folds or weak denticles. Lateral teeth
long, broad.
Remarks.
Thais (Mancinella) echinata (Blainville, 1833) differs
pamarily in having a more rounded shell with 4 spiral
rows of short, acute spines on the last teleoconch
whorl:, its aperture 1s relatively smaller with more
numerous, weaker lirae and broader columellar lip,
with a less well delimited edge. The Japanese species
T. siro (Kuroda, 1931) has 4 spiral rows of short
spines, a more indistinct edge of columellar lip, more
flattened spiral cords on the last teleoconch whorl, and
4 low, weakly elongate nodes within the outer apertural
edge instead of 8-10 lirae. At first sight, the shell also
resembles Drupella cornus (Rôding, 1798) and D.
eburnea (Küster, 1862), but both species are narrower,
have fewer, broader spiral cords, a denticulate instead
of lirate outer apertural lip, a narrower aperture, and a
higher spire. Morcover, the radula of Drupella is quite
different from that of Zhaïs, and from other rapanines,
in having long and slender, reed-shaped laterals.
Etymology.
Named for Dr. Dai Herbert (Natal Museum,
Pietermaritzburg), known for his numerous papers on
South African Trochoidea, and who took the beautiful
photograph of 7° herberti in situ.
ACKNOWLEDGEMENTS. This study was made possible
thanks to the helpful collaboration of many people. For
the material Ï am thankful to P. Bouchet (Muséum
National d'Histoire Naturelle, Paris); P. Clarckson (Port
Lincoln, South Australia); A. Deynzer (Sanibel,
Florida), A. Edinger (Mandurah, Western Australia):
R.N. Kilburn (Natal Museum, Pietermaritzburg), and I.
Loch and W.F. Ponder (Australian Museum, Sydney).
Many thanks also to K. Hasegawa (Showa Memorial
Institute, Tsukuba, Japan), and to T. Okutani (Nihon
University, Kanagawa-ken, Japan) for helpful
information and for the loan of Kuroda's type
specimens. | am also much indebted to A. Warén
(Naturhistoriska Museet, Stockholm, Sweden) for
processing and SEM of the radulae, and to B. A.
Marshall (Museum of New Zealand, Wellington) and
EH. Vokes (Tulane University) for their constructive
comments, suggestions, and linguistic assistance.
HOUART Eight new species of Muricidae APEX 13(3): 95-109, 20 juil. 1998
REFERENCES
HOUART, R. 1986. Mollusca Gastropoda: Noteworthy
Muricidae from the Pacific Ocean, with description
of seven new species. Résultats des campagnes
Musorstom I & II, Philippines. Mém. Mus. natn. Hist.
nat. sér. a. 133: 427-455.
HOUART, R. 1993. Description of three new species
and one new subspecies of Muricidae (Muricinae and
Muricopsinae) from West Africa. Boll. Malac. 29(1-
4): 17-30.
HOUART, R. 1997. Mollusca, Gastropoda: The
Muricidae collected during the KARUBAR Cruise in
eastern Indonesia. Résultats des Campagnes
MUSORSTOM, 16. Mém. Mus. natn. Hist. nat. 172:
287-294.
Lussi, M. & D. BRINK. 1996. A collectable group for
all -Thaïis and allied genera in South Africa. The
Strandloper 248: 1, 3, 6-7, 12.
TRÔNDLE, J. & R. HOUART. 1992. Les Muricidae de
Polynésie Française. Apex 7(3-4): 67-149.
Fig. 37. Ergalatax dattilioi n.sp. Japan, Wakayame Pref., Minabe, 100-120 m, coll. R. Houart, 17.7 mm. Figs 38-39.
Ergalatax tokugawai Kuroda & Habe, 1971. Japan, Miura Peninsula, Kanagawa Pref., WS, 2 km off Jôgashima, 78-
85 m, holotype NSMT-MOR 9604, 16 mm.
107
APEX 13(3): 95-109, 20 juil. 1998 Eight new species of Muricidae HOUART
Figs 40-41. Radulae (scale bar: 50 um). Fig. 40. Attiliosa edingeri n.sp. West Australia, off Esperance. Fig. 41.
Thais (Mancinella) herberti n.sp. South Africa, Zululand. Fig. 42. Thais (Mancinella) herberti n.sp. /n situ, on
Sponges, paratype NM E8760/T1470, photo D. Herbert. Figs 43-44. Protoconchs (Scale bar: 0.5 mm). Fig. 43.
Orania rosadoi n.sp. Fig. 44. Ergalatax dattilioi n.sp.
108
HOUARI Eight new species of Muricidae APEX 13(3): 95-109, 20 juil. 1998
30
30
45
15 30 45 60 75 90 105 120 135 150 165 180 165 150
Figs 45-47. Distribution maps Fig. 45. Ergalatax dattilioi n sp. Fig. 46. Apixystus rippingalei n.sp. Fig. 47
Trophonopsis bassetti n.sp.
109
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Revision of the F'asticardium assimilée Species group
Taxonomic revision of the Indo-Pacific
Vasticardium assimile species group (Mollusca, Cardiidae)
Jacques VIDAL
Attaché au Museum National d'Histoire Naturelle de Paris,
Laboratoire de Malacologie, 55 rue Buffon, 75005 Paris, France
KEY WORDS. Mollusca. Bivalvia. l'asticardium. Indo-Pacific.
SUMMARY. Several forms of Indo-Pacific l'asticardium of medium size. are here grouped as the
Vasticardium assimile species group. Threc of these forms. mainly from the Indian Ocean and
previously described as threc different species. 1° assimile. 1° lacunosum and 1° rubicundum. share
many characters and have often been confused. They are considered here to represent only two
species, }° assimile and T° rubicundum. While 1° assimile lacunosum is treated as a subspecies
restricted to the Persian Gulf. } assimile is restricted to the western Indian Ocean, southernmost Red
Sea and Persian Gulf, whereas 1° rubicundum, although more abundant in the southeastern Indian
Ocean, extends also into a large part of the western Pacific, where it has been described as
mindanense, compunctum., and kengaluorum. Two rare additional species, Fasticardium rhegminum.
from the Masirah area (Oman). and l’asticardium thomassini sp. nov. from southeastern Africa and
APEX 13635 111-125. 20 juil 199$
Madagascar, are treated.
MATERIAL, METHODS AND TEXT
CONVENTIONS
The material comes from the following museums:
AMS: Australian Museum, Sydney.
ANSP: Academy of Natural Sciences, Philadelphia.
BMNH': The National History Museum, London.
IRSNB: Institut Royal des Sciences Naturelles de
Belgique, Brussels.
LACM: Los Angeles County Museum of Natural
History, Los Angeles.
MHNG: Museum d'Histoire Naturelle de Genève,
Geneva.
MNAN: Museum National d'Histoire Naturelle, Paris.
NHMVW: Naturhistorisches Museum, Vienna.
NMVW: National Museum of Wales, Cardiff.
RMNH: Nationaal Natuurhistorisch Museum, Leiden.
USNM: National Museum of Natural History.
Washington.
ZMA: Zoologisch Museum. Amsterdam.
ZMUC: Zoologisk Museum, Copenhagen.
In the description of the shells, particular attention was
devoted to the rib morphology (structure and
ornamentation of the ribs), this in an ontogenic
perspective, by taking into account these elements in
the earliest parts of the shell and their variations
towards the adult parts, in the various "quarters".
For examination purposes, the shells are divided
externally into four radial "quarters": AQ: Anterior
Quarter, MAQ: Medio-Anterior Quarter. MPQ:
Medio-Posterior Quarter: PQ: Posterior Quarter.
Longitudinally. shells are divided schematically into
two parts, a "juvenile" (or umbonal) part and an "adult"
(or marginal) part. In the description of the hinges,
angle A 1s the angle formed by the two lines joining the
tip of the main cardinal tooth to the ups of the laterals
respectively. Ratio D is determined by dividing the
length of the line from the tip of the umbo to the tip of
the posterior lateral by the corresponding distance from
the umbo to the tip of the anterior lateral.
SYSTEMATICS
Family CARDIIDAE Lamarck, 1809
Genus Vasticardium Iredale, 1927.
Type species: Cardium elongatum Bruguière, 1789, by
original designation., Iredale 1927: 75.
Diagnosis. Shells medium to large, generally elongated
and variably inflated. Shape ovoid and symmetrical or
posteriorly expanded, obliquely or not. “winged" or
truncated. Mean rib number ranges from 30-40,
exceptionally 45. Hinge line moderately angled (Angle
A circa 130°). Cardinal teeth in right valve separated or
merely touching at their base and never connected by a
high and narrow dorsal saddle.
In juvenile median and anterior parts, ribs quickly
become high. square-sided and fully ornamented,
directly following millimetric smooth, very primitive
shell. In PQ, nbs always high and square-sided, always
simple, not longitudinally divided into two parts; top
scales or nodules always arranged in a single row along
apex. In other quarters of adult shells, rnbs generally
high. often squared and overhanging interstices, rarelv
111
APex 1303) 111-125, 20 juil 1998
Revision of the l'asticardinnr assimilé species group
VIDAI
triangular, in MPQ they often bear scales or tubercles
with crenulated margins, cross-bars in the anterior half
Interstices moderately deep and wide, with à smooth or
fincly striated flat bottom, independent from flanks of
nbs
Remark. In previous papers (VIDAI. 1992, 1993), 1
provisionally used Acrosterigma Dall, 1900, for specics
belonging to the same genus as the one described here
l currently consider the genus l'asticardium, as defined
above, to be more appropriate
DIAGNOSIS OF THE VFASTICARDIUM
ASSIMILE SPECIES GROUP
Species of the group. all of medium size, share several
similar macroscopic characters such as shape. hinge
structure, nb number, and above all color. In addition.
the young shells (up to one or two cm in height or
shightly more) are somewhat similar as far as nb
sculpture is concerned: in PQ, rather low, asymmetrical
nbs bearing top oblique scales. with à sharp anterior
margin overhanging interstices (Fig. 8) and, in other
parts of shell, high nbs with à roundly triangular top
zone crenulated on margins which often overhang
smooth and flat-bottomed interstices (Figs 6-7).
Another significant similanty 1s that the carbonates
which form the shell seem to have a characteristic thin
microscopic structure which makes the shell fragile.
and easily faded. worn or notched, and gives it
lustreless colorations. In addition, nbs are constituted
of several carbonaceous laminae, longitudinally
jJuxtaposed, some of them with pink coloration visible
when nbs are worn.
Vasticardium assimile (Reeve. 1844)
Figs 1-11, 25-26.
Cardium assimile Reeve, 1844, Sp. 45, pl. 9, fig. 45.
Cardium lacunosum Reeve, 1845. Sp. 81, pl. 16, fig.
81.
Selected references.
Cardium assimile: Rômer, 1869, pl. 11, fig. 11.
Cardium assimile: Braga, 1952: 49, pl. 10, fig. 2.
Laevicardium (Trachycardium) rubicundum:
Fischer-Piette, 1977: 65 [Partly].
Trachycardium lacunosum: Bosch, 1982: 172.
lrachycardium lacunosunr. Smythe, 1982: 100, pl.
18. fig. c
Trachvcardium elongatunr: Drivas & Jay. 1988: 140,
pl. 55. fig, 1
Trachvcardium lacunosumr: Oliver, 1992: 126, pl. 23.
fig. 8a-b.
Acrosterigma assimile: Oliver, 1995: 246, fig. 1091.
NOT 7rachvcardium lacunosum: Keen, 1945: 36 [-
probably rubicundum].
Type material.
Cardium assimile : Three syntypes in BMNH Reg. N°
1978-129, from Zanzibar. The largest 1s figured here
(Figs 1-3). Another specimen corresponds exactly to
the dimensions of Reevc's figure (H= 56.0, L= 42.7
min). but looks different because the illustration 1s of
poor quality.
Cardium lacunosum: Once shell considered as
holotype in BMNH Reg. N° 1978-133 locality
unknown (Figs 4-5). The right valve fits the figured
one as far as dimensions and rib number are concerned,
but several other characters do not match Reeve's
figure and description. For example, nibs are curved
backwards in projection with thin numerous marginal
crenulations (at least 40: sec definition p. 11-2) in the
supposed "holotype". as opposed to perfectly straight
nbs with strong rarer crenulations (barely 20 in
number) in Reeve's specimen. For these reasons I think
that the shell in the BMNH, which is a "common" form
of F° assimile, is not the shell described and figured by
Reeve as lacunosum [which means "hollowed", an
allusion to the strong marginal crenulations of the nbs;
Reeves writes "laterally hollowed"], and a neotype will
be selected for these special forms from the Persian
Gulf considered as a subspecies: F. assimile lacunosum
(see below).
Description.
Shells medium to large. Shape regularly subovoid,
almost equilateral but somewhat posteriorly truncated,
with ribs sometimes slightly curved backwards in
projection. Generally elongated in the adult stage
(mean L/H= 0.77: range 071-092), and relatively
tumid (mean W/L= 0.87; range 079-102).
Lunule narrow but well marked, and purple in color.
V. assimile 1s always externally colored with stripes
and/or splashes of vivid but lusterless brown or purple,
more rarely orange or pink; interior white, sometimes
with a double colored ray in umbonal area, and purple
margin.
Figs 1-3. Vasticardium assimile, Syntype, BMNH 1978-129, L= 470 mm. Figs 4-5. Vasticardium assimile,
specimen erroneously considered as holotype of Cardium lacunosum Reeve, BMNH 1978-133, L= 40.5 mm. Figs
6-7. Vasticardium assimile, specimen from Magaruque Is, Mozambique, MNHN. Fig. 6: left valve; detail of juvenile
median part of shell, scale x 5. Fig. 7: right valve, detail of juvenile PQ and MPQ; scale x 5. Fig. 8. Vasticardium
assimile, right valve, specimen from Zanzibar, MNHN, detail of juvenile MPQ and MAQ, particularly thin and
numerous marginal crenulations on the ribs (50 per two cm), scale x 25.
112
VIDAL Revision of the Vasticardium assimile species group APEX 13(3): 111-125, 20 juil. 1998
113
APEX 13(3} 111-125, 20 juil. 1998
Revision of the Fasticardium assimile species group
Hinge moderately arched (mean < A= 130°, range
120-135°) and rather asymmetrical (mean ratio D-
1.20, range 100-145). Posterior cardinal tooth in left
valve long (2/3 of the width of the linge area) and high
(almost as high as main cardinal), relauvely narrow and
often with an acute summit. Foundation of anterior
lateral (mainly in left valve, tooth PIT) very moderately
“hook-shaped". A medial short weak nb (umbonal
support) present in umbonal cavity of many shells.
Mean rib number 33.4 (range 31-37).
Rib morphology:
In juvenile PQ (Fig. 8) nbs rather low and
asymmetrical, with anterior edge overhanging
interstice, with a sharp continuous margin jutted in
front of scales. Rib tops set with large, more or less
tubercular, slightly twisted oblique scales: no
secondary small scales occur on edges of ribs as in
some other species groups of l’asticardium: interstices
as wide as nibs. In adult PQ, scales on nb tops can
become more or less rounded tubercles, often irregular
in shape and degenerate.
The juvenile median part of shell (Fig. 6) bears high
nibs, of trapezoidal section with wide sloping flanks
and a slightly rounded smooth top zone, bordered on
both sides by crenulations which overhang the flanks:
tops sometimes regularly swollen by extension of
marginal crenulations. Interstices also trapezoidal in
section, and almost as large as nbs; their bottoms are
flat and clearly separated from flanks of nbs by a break
line. This bottom often transversely very finely notched
and flanks sometimes distinctly grooved, mainly
posteriorly. In adults, a significant change from the
juvenile sculpture occurs in MPQ (Fig. 25): the last nbs
of this quarter (close to PQ) become very asymmetrical
with anterior flank shorter and more abrupt; on anterior
margins of nbs, the crenulated projecting edge does not
change, but on the wider posterior side, marginal
crenulations progressively join the flank ridgelets
which become stronger, and overhanging disappears.
Posterior parts of ribs form a wide dipping flank,
entirely strongly ridged, ridges and furrows becoming
more and more irregular with age; this evolution
(illustrated in Fig. 25) is very characteristic of
Vasticardium assimile and examining it always allows
to separate this species. The reduced nb tops remain
smooth or become regularly ridged in the most adult
part, rarely with a herringbone structure. In anterior
part of adult MPQ and MAQ, ribs remain crenulated on
both overhanging margins.
Figs 9-11.
VIDAL
In juvenile AQ, nibs more square-sided, and slightly
asymmetrical, with transverse ridges on their top (Fig.
6). interstices rounded, without any discontinuity with
nb flanks. both being slightly striated by fine growth
lines. In adult AQ, tops of ribs become flat and
depressed antériorly and are ornamented with shightly
curved, strong imbricated cross-bars.
Material examined.
The following lots in addition to the type material:
A) South Africa: East London: I(MNHN);:
Sheptone: I(MNHN), I(ANSP)
B) Mozambique: Magaruque Island: I(MNHN),; Santa
Port
Carolina Island: 2(MNHN). no specific location:
I(MNHN)
C) Madagascar: Tulear: 2(MNHN), no specific
location: IIMNHN). I(BMNH)
D) TANZANIA: Zanzibar: 5S(MNHN), 2(BMNH),
I(ANSP), I(USNM). I(LACM), M'Boa Mai:
I(USNM)
E) Kenya: Shimoni: I(BMNH),; no specific location:
I(MNHN)
F) Somalia: Eil: I(MNHN): Beidi: I(MNHN);:
Mogadishu: IIMNHN), I(ANSP), S. Cape Guardafui:
I(MNHN); Bender Kassim: I(MNHN). W. Elayu:
I(MNHN), E. Sugra: I(MNHN), Mait I(MNHN);
Candara: I(ANSP)
G) Aden: I(MNHN), 1I(BMNH)
H) Djibouti: 2IMNHN)
I) Yemen: Abd el Kun Island: I(MNHN); Socotra
Island: I(MNHN): Mocha: I(MNHN);, Hodeida:
2(MNHN), 2(USNM); N. Hodeida: I(MNHN)
J) Oman: Masirah: I(MNHN), Musqat: I(MNHN);
I(BMNH)
K) Persian Gulf: Bender Abbas: I(MNHN); Khassab:
I(BMNH), Trucial Coast: 2(BMNH), Lavan Is:
I(MNHN), AI Hamra: I(MNHN), Doha: I(MNHN):
Dammam: I(USNM);, Ras Tannurah: 2(ANSP),
I(USNM), Kuwait, AI Fintas: I(MNHN), Kuwait:
I(USNM). not localized: Tarut Bay: I(ANSP),
I(USNM), Chaschuse Is I(USNM); no specific
location: I(BMNH); 1(LACM)
L) Pakistan: Karachi: I(BMNH), I(USNM)
M) India: Tuticorin: I(MNHN), Gulf of Manaar:
2(ANSP), I(BMNH)
N) Sri Lanka, Trincomalee: 1 (BMNH); no specific
location: 2(BMNH); 2(USNM); 1(LACM)
O) No locality: 3(MNHN).
Vasticardium assimile lacunosum, Neotype, MNHN, ex coll. Vidal, L= 460 mm. Figs 12-14.
Vasticardium rubicundum, Syntype, BMNH ex coll. Cuming, L= 49.0 mm. Figs 15-17. Vasticardum rhegminum,
bivalved specimen, MNHN ex coll. Sue Hobbs, L= 39.4 mm. Figs 18-19. Vasticardium rubicundum, Lectotype of
Cardium mindanense, BMNH 1070-124, L= 34.7 mm. Fig. 20. Vasticardium rubicundum, Paralectotype of Cardium
mindanense, BMNH 1070-124, L= 340 mm. Figs 21-24. Vasticardium thomassini, Holotype, MNHN ex coll.
Thomassin, L= 27.7 mm.
114
VIDAL Revision of the Vasticardium assimile species group APEX 13(3}) 111-125, 20 juil. 1998
fl At *
TRE At 134 #3
Haut LAN AS
* à
115
APEX 13(3): 111-125, 20 juil. 1998
Revision of the Fasricardium assimile species group
VIDAL
Measurements of !° assimile ss, excluding Persian Gulf forms.
Height Length Width L/H W/L ratio D | angle A
[ (mm) (mm) (mm) £
Syntype 633 47.0 40.0 0.74 0.85 LIST OT
| Syntype 56.0 42.7 353 0.76 0.83 1.20
Mozambique 17 40.0 34.4 0.74 0.86 127
| Madagascar 57.0 47.0 41.3 0.82 0.88 1.05
| Zanzibar 59.4 443 37.2 0.75 0.84 1.15 120
Kenya 474 355 306 | 075 | 08 1250 [T2
Socotra Is. 532 34.0 0.77 0.84 PA TONNES
N Somalia 42.1 0.93 1.17 125
ie 440 293 081 0.82 131
Djibouti 39.0 246 0.79 0.79 1.15
N Yemen 48.9 34.5 2221 0.71 0.93 1.26
| Masirah Is 558 438 364 0.78 0.83 1 09
S India 449 36.5 31.2 0.81 0.85 1.60
The maximum size observed in the "nominal" form is a right valve from Madagascar, in MNHN (H= 68.8 mm, L=
49.0 mm).
Distribution.
Because of the possible confusion with F. rubicundum.
it is impossible to accept literature records uncritically.
From material examined, I have ascertained the
following distribution: east coast of Africa from East
London (Natal) to the Gulf of Aden, Madagascar,
Seychelles, southernmost part of the Red Sea (Yemen),
Oman (Masirah, Muscat). Persian Gulf (from Strait of
Hormuz as far as Kuwait). Pakistan (Karachi). south
India (Gulf of Mannaar). and Sri Lanka.
It is also present in Réunion and Mauritius (DRIVAS
& JAY, 1988, pl. 55, fig. 1, under the name e/ongatum).
The only record in the literature of F! assimile outside
these areas 1s from Java (ALTENA, 1945: 150). There
are records of F° /acunosum from Torres Strait
(MELVILL & STANDEN. 1899: 190) and New Guinea
(KEEN, 1945: 36). I consider all these records to
probably refer to l! rubicundum which has à western
Pacific extension.
Remarks.
Differences from F° rubicundum: V. assimile 1s very
close to }! rubicundum, mainly in the juvenile shells.
For differences see under }. rubicundum.
Variability of F! assimile: This species has many
constant characters always allowing one to separate it
from the others, both within the present group and
outside it [It is. particularly, easily separable from
Acrosterigma magnum (Linné), although it 15
considered by Reeve "quite similar" (= assimile) to this
species]. Nevertheless some characters can vary
according to populations and/or individuals. The most
significant of these variations are:
1- Elongation in adult specimens (L/H range 0.71-
116
0.92).
2- Size and number of the marginal crenulations of
the ribs: These are very variable mainly in the median
part of the shell, from very small and reduced to thin
scales [in which case the ribs in the median part remain
roughly triangular and the interstices opened and wide
(Figs 8, 25)], to as wide as about half the width of the
nb [in which cases the crenulations overhang the flanks
of the ribs which are nearly "vertical", and the
interstices are reduced in width (Fig. 26)]. Apparently
in relation to the size, the number of the crenulations 1s
also variable: in the middle of the shell, in a band
between 3 and 5 cm from the tip of the umbo (curved
distance). this number varies from 21 to 52.
3- Regular swelling or ridging of the tops of the nbs
in the median part of the shell: In the anterior part of
the shell, the rib tops always bear cross bars jJoining the
lateral crenulations, this phenomenon gradually
disappears backwards and, in the medial part of the
shell. the tops can either still be swollen or become flat
and smooth. This character varies also with the age of
the shell. and the swelling of the tops 1s variable
according to individuals, the young shells developing it
relatively late.
4- Shape and size of the top ornaments of the nibs in
PQ: These ornaments can vary in the same individual,
and also become locally irregular and degenerate: they
vary from nodular or elongated tubercles to thin,
variably twisted, oblique lamellar scales.
These variations generally occur at random and
separately according to populations and/or individuals.
However a geographical group with constant
simultaneous variations is present in the Persian Gulf,
.and is here defined as a subspecies: 1° assimile
lacunosum.
VIDAL
Vasticardium assimile lacunosum
(Reeve, 1845)
Figs 9-11, 26.
Cardium lacunosum Reeve. 1845, Sp. 81, pl. 16, fig.
81.
Acrosterigma lacunosa: Oliver, 1995: 246, fig. 1090.
Type material.
As seen above, the specimen BMNH 1978-133 1s
closer to the nominal subspecies of |’! assimile and 1s
certainly not the holotype of C. /acunosum figured and
described by Reeve. Considering that this latter shell 1s
not traced, a neotype of Cardium lacunosum 1s selected
here (Figs 9-11), a shell from AI Fintas, Kuwait, in
MNAN, measuring 57.7 x 46.0 x 41.3 mm, with 33
ribs. Ratio D= 1.27, < A= 130°: number of crenulations
27.
Revision of the Vasticardium assimile species group
APEX 13(3): 111-125, 20 juil. 1998
Description.
Shells medium to large. Shape regularly subovoid,
almost equilateral but somewhat posteriorly truncated,
with ribs sometimes slightly curved backwards in
projection. Generally moderately elongated in adult
stage (mean L/H= 0.81; range 0.73-0.89), and
relatively tumid (mean W/L= 0.90; range 0.84-1.02).
Lunule, colour, hinge, rib number in agreement with
general description of species. No umbonal support
observed.
Rib morphology:
In PQ, thin, lamellar, homogeneous and regularly set
oblique scales on top of ribs (Fig. 26).
In median and anterior parts, few and strong
marginal crenulations of ribs, and lack of smoothness
of ribs tops.
Material examined and distribution.
Persian gulf (see above).
Measurements of l. assimile lacunosum
Height
(mm)
ratio D
Bandar Abbas
W/L
46.0 0.80 127 E
Kuwait 58.1 0.76 0.98 130
0.73
26 55
1.02
Maximum record in literature, a height of 90 mm (Smythe, 1982).
Remarks.
The uniqueness of the subspecies /acunosum 1s the
constant conjunction of two characters which can
sporadically exist in the nominal subspecies:
1- Thin, lamellar, homogeneous and regularly set
transverse scales on top of nibs in PQ (Fig. 26), rather
than the globular, somewhat elongated variable
tubercles which are generally present in the nominal
subspecies (where comparable lamellar scales can also
sporadically exist, Figs 7, 25).
2- Strong marginal crenulations of the nbs, and lack
of smoothness of rib tops: In the median part of the
shell] the lateral crenulations proceed up to the top zone,
forming cross-bars or partially joined rugae, all of
which gives the ribs a "“plaited-like structure" as
Smythe says (1982: 100), figuring a good example of
this special form (1982, pl. 18, fig. c). This character
already appears in very young shells: in the umbonal
area the crenulations of the nbs become regularly
swollen very early, unlike those in the nominal
subspecies. The number of crenulations is smaller: 21
to 37 crenulations (as defined above) versus 35 to 52 in
the nominal subspecies.
Besides the two fundamental elements of distinction
above, some other more statistical elements seem to
characterize lacunosum. It is less elongated (average
L/H= 0.81, vs general average= 0.77), more tumid
(average W/L= 0.90, vs general average= 0.87), and
larger (in the largest specimen H= 90 mm, as opposed
to 68.8).
Vasticardium rubicundum
(Reeve 1844.)
Figs 12-14, 18-20, 27, 31.
Cardium rubicundum Reeve, 1844, Sp. 44, pl. 9, fig.
44.
Cardium mindanense Reeve, 1844, Sp. 19 [excluding
the syntype figured pl. 9, fig. 19 = 7rachycardium
egmontianum Shuttle-worth, 1856, 5: 172].
lasticardium compunctum Kira, 1959: 137, pl. 55.
fig. 9.
Acrosterigma kengaluorum Voskuil & Onverwagt
1992: 33. pls 1-2.
17
APEX 1303) 111-125, 20 juul 1998
Revision of the P'asricardiunm assinnle species group
VibaAI
Selected references.
l'asticardium rubicundum: Keen. 1945: 36
Cardium rubicundunr Braga. 1952: 49, pl 10, fig. 2
NOT Laevicardium (Trachvcardium) rubicundum
Fischer-Piette, 1977: 65 [= pantly l'asticardium
assimile]
lrachvcardium rubicundum: Oliver, 1992: 126, pl
23, Figs 7a-b
lrachvcardium (l'asticardium) mindanense
Spangsteen & Leobrera. 1986: 306, pl. 87, fig. 2
Type material.
Cardium rubicundum : Three syntypes from Zanzibar
in BMNH., coll. Cuming. all larger than Reeve's figure:
the largest (Figs 12-14) measures 60.0 X 490 x 41,3
mm
Cardium mindanense : The name of this taxon is not
to be confused with C mendanaense Sowerby. 1897.
which 1s another Indo-Pacific shell belonging to the
group of l'asticardium orbita (see VIDAL 1997a).
There are three syntyvpes of Cardium mindanense
Reeve in BMNH (ex Cuming) said to have come from
Mindanao (Philippines). As indicated by pencilled
notes, these three specimens were initially set on the
same board. They are now separated into two boxes :
- one box with the specimen figured by Reevc,
subsequently erroneously labelled "holotype". Reg. N°
1978-124, measuring 41.0 x 35.2 x 29.0 mm, with 30
nibs. This shell 1s 7rachyvcardium egmontianum
Shuttleworth, 1850, from the Atlantic coast of North
America. This shell was probably placed with the
others by Cuming. in error, because of a certain
superficial resemblance between them. Further. it was
probably chosen by Reeve for the figure because it was
the largest.
- a second box with two specimens of J°
rubicundum, respective dimensions: H= 40.1 mm, 37.6
mm; L= 347 mm, 340 mm; W= 30.6 mm. 253 mm:
the number of ribs is 36 in both specimens. These last
two shells are very probably those which come from
Mindanao and from which the name mindanense
originates. In order to avoid possible confusion and to
preserve the legitimate Indo-Pacific origin of thus
taxon, [| have selected as lectotype of Cardium
mindanense the largest of these (Figs 18-19): the
remaining syntype. becomes the paralectotype (Fig.
20).
Acrosterigma kengaluorum:. The holotype is in
RMNH (56769) from Homara. Guadalcanal Is. the
Solomons., à shell measuring 30.7 x 27.8 x 21.0 mm.
with 34 nbs Twenty paratypes. all in private
collections. are from the same area; their dimensions
arc as follows: H= 24.1 10 37.4 mim. L= 196 to 345
num. W= 152 to 256 mm Mean rib number 35.4
(range 32-38) Mean L/H= 090 (range 086-096).
mean W/L= 0.75 (range 0.69-0.82).
l'asticardium compunctum. This name first appears
as a Kuroda MS. name in HABE (1951: 145). KIRA
(1955) gives à figure and sull credits the name to
Kuroda MS. but this figure 1s not accompanied by à
description. The name 15 first validated by KIRA (1959:
139) who gives a diagnosis of the species in Japanese.
using the same figure. In 1962 (p. 156). KIRA gives a
description in English. The shell figured by KIRA (1955
and 1959, pl. 55, fig. 9, and 1962, pl. 56. fig. 9) is
considered as the holotype. with dimensions of 41.5 x
32.9 OX 315 mm. According to a personal
communication from À. Matsukuma. Kira's 1llustrated
material is stored in Osaka City Museum (Natural
History) and 15 not available for loan. The localities for
the species given by KIRA, in 1962, are the Amami and
the Ryukvyu Islands (Japan). Only photographs of Kira's
type of !° compunctum have been examined but one lot
in MNHN comes from the type locality, the Amami
Islands. Japan.
Description.
Shells regularly subovoid, almost ellipsoidal in shape
and almost equilateral. except for à small truncation in
posterior margin. PQ is somewhat flat or slightly
depressed and forms an obtuse angle with rest of shell.
Ribs generally straight, rarely shghtly curved
backwards in projection, adult state little elongated
(mean L/H= 981: range 077-086) and relatively
tumid (mean W/L= 0.84: range 0.78-0.84).
Exterior splashed or striped with red-purple. or
sometimes brown; yellow or light orange markings can
occur and predominate in specimens from southern
Japan, and brown markings predominate in rest of
Pacific: entirely white specimens are rare. Interior
white with a colored margin and an umbonal area with
two rays. Lunular area narrow, generally purple
colored.
Fig. 25. Vasticardium assimile, left valve, specimen from Magaruque Is, Mozambique, MNHN (same as Figs 6 and
7), detail of PQ and MPQ showing particularly the ontogenic change towards the shell margin of the small posterior
marginal crenulations into elongated ridges on the flank of the ribs in last ribs of MPQ, scale x 25 Fig. 26.
Vasticardium assimile lacunosum, left valve, specimen from the Persian Gulf, detail of PQ and MPQ; scale x 25.
Fig. 27. Vasticardium rubicundum, left Valve, specimen from Zanzibar, MNHN, detail of PQ and MPQ showing
particularly the additional scales in MPQ, superposed on the thin marginal crenulations, scale x 2.5. Fig. 28.
Vasticardium thomassini, Holotype; detail of PQ and MPQ, scale x 4. Fig. 29. Vasticardum rhegminum left Valve,
same specimen as in Figs 15-17, detail of PQ and MPQ, scale x 25 Fig. 30. Vasticardum rhegminum, small right
valve, MNHN coll Day, detail of juvenile PQ and MPQ showing particularly in PQ the anterior edge of ribs with a
thin longitudinal furrow, scale x 5
118
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APEX 13(3): 111-125, 20 juil. 1998
Revision of the Vasticardium assimile species group
VIDAL
119
APEX 13(3} 111-125, 20 juil. 1998
Hinge moderately arched (< A range 125-140°) and
rather asymmetncal (mean ratio D= 1.15; range 1.0-
1.43). Postenor cardinal in left valve (4b) rather long
and high, sometimes with à sharp top; foundation of
anterior laterals, mainly in night valve, moderately
projecting and more hook-shaped than in F assimile
No medial nb in umbonal cavity.
Mean rib number 36.0 (range 33-39).
Rib morphology:
In juvenile PQ, nbs rather low, and asymmetrical,
with anterior edge overhanging interstice with a sharp
continuous rib margin jutted in front of scales or rarely
dividing into successive transverse bent lamellae; rib
tops bear large somewhat tubercular, slightly twisted
scales, obliquely set; interstices are as wide as nbs. In
adult PQ, anterior edge of nbs generally becomes less
sharp and often does not overhang interstices which
become narrower than nbs; scales become tubercular
(Fig. 27).
In juvenile median part of shell, ribs trapezoidal with
rounded, narrow, almost flat tops, which are always
smooth, flanks of ribs strongly ridged and furrowed,
with furrows generally straight except for anterior
flanks of MPQ, where they are very bent. These
ornaments disappear near base of flanks, leaving a
smooth basal zone. Before disappearing, these ridges
thicken, forming small tubercles on their lower
extremity, these tubercles, being always aligned at
lower 1/3 of ribs, form regular beaded longitudinal
lines along flanks of ribs; these "beads" extend into
interstices, sometimes almost touching, in that latter
case, the smooth inferior part of ribs and bottom of
interstices form "tunnels." Bottom of interstices flat
and weakly notched. Margin of top zone very variable.
with respect to crenulations, from one population to
another, these crenulations may or may not be present.
on one or on both sides of ribs, but in anterior half of
shell crenulations are always in continuity with the
flank ndges which are strongly furrowed but not
beaded. When these crenulations are present. flanks
bear two longitudinal lines of tubercles: crenulations in
the upper part, and "beads" in lower part. In adult
MPQ, ribs remain or become very asymmetrical: on
anterior part of nibs the finely crenulated projecting
edge remains the same, but furrowing of the reduced
flank lessens; in the wider posterior part. crenulations
and flank furrows progressively disappear, and strong
oblique straight scales develop on top and flank of rib.
These scales, which are a continuation of main scales
of adjacent PQ. superpose on pre-existing
Revision of the Fasticardium assimile species group
VIDAL
ornamentation and entirely replace it in the most aged
part of shell (Fig. 27). In adult MAQ, and sometimes as
early as in anterior part of MPQ, nbs become very
asymmetrical, upper anterior part of rib flanks lowers
forming à gently sloping ridged plane with the anterior
part of top zone, long ridges of these planes form
typical asymmetrical herringbone patterns with
posterior crenulations or ndges, these herringbone-
shaped ridges, first open then closed on all ribs, are
often of a light color and contrast strongly with the
darker color of nbs.
In juvenile AQ, flanks of ribs not beaded, interstices
only finely striated by growth lines, and crenulations
tend to join to form imbricated transverse ridges with a
quasi-herringbone pattern. In adult AQ, these patterns
evolve into imbricated continuous cross bars on top and
anterior flank of ribs, tops retain a generally
asymmetrical hernngbone aspect.
Material examined.
Type material: Syntypes of C. rubicundum and C
mindanense, Holotype and paratype N° 1 of À.
kengaluorum, and the following lots:
A) South Africa: East London: I(MNHN), I(USNM);
Port Sheptone: I(MNHN) I(ANSP);, S. Durban:
I(MNAN), ON. Durban I(MNHN), Brighton:
I(MNAN), no specific location: 1I(BMNH)
B) Mozambique: Joao Belo: I(MNHN), Inhambane:
I(MNHN), Chidenguele: I(MNHN), Ponta da Barra:
I(USNM)
C) Madagascar: Tulear: S(MNHN)
D) Tanzania: Dar es Salaam:
4(MNHN). I(BMNH), 2(ANSP)
E) Kenya: Shimoni: 4(BMNH)
F) Somalia: N. Mogadishu: I(IMNHN), I(ANSP)
G) "Indian Seas": IIMNHN)
H) Sri Lanka: no specific location: I(IMNHN,) I(AMS)
1) Singapore: I(AMS)
I(AMS), Zanzibar:
J) Indonesia-Java: Batavia Bay: 1(ZMA). Bantan:
2(USNM)
K) Indonesia-Bali: Nusa-Dua 1(LACM)
L) Philippines: Mindanao, Zamboanga: I(MNHN):
Luzon,Tabangao: I(AMS): Luzon. Corregidor:
I(AMS)
M) Indonesia-Sulawesi: N. Paleleh: 1I(RMNH)
N) Indonesia-Moluccas: no specific location: 1(ZMA)
O) Papua New Guinea: Oro Bay: I(MNHN)
P) Japan: Amann Islands: I(MNHN).
Okinawa: 2(USNM). I(LACM).
I(ANSP).:
Fig. 31. Vasticardium rubicundum, left valve, specimen from Zanzibar, MNHN, detail of median part (MPQ and
MAQ). Scale x 35. Fig. 32. Vasticardium thomassini, left valve, holotype, detail of median part (MPQ and MAQ),
scale x 4.0.
120
VIDAL
Revision of the Vasticardium assimile species group
Measurements of }. rubicundum
(mm)
a
[a
0.84 1.30
Syntype
0.84
Syntype
Zanzibar
Indian Seas
Sri Lanka
Japan, Amami
1.18
0.82 1.24
0.88
HT RES
EE 81 0. Re IE 17 1350
0.86 0.81 1.43 1400
Largest specimen observed 1s the above syntype (H= 60.0 mm).
Distribution.
Because of the frequent confusion of 1° rubicundum
with F assimile, the distribution derived from the
literature 1s unreliable. and here only the verified
localities are taken into account. Its extension into the
western Indian ocean is smaller than the one of J:
assimile, and it is not recorded from the Gulf of Aden
or Arabian Sea. But J° rubicundum has a larger Pacific
distribution. and 1s sporadically found as far as in Japan
and the Solomons. However. it is undoubtedly locally
rarer outside the southwestern zone of the Indian Ocean
where it is sympatric with }° assimile.
Remarks.
Variability of 1° rubicundum: Unhike 1: assimile, V
rubicundum is stable in shape and rib ornamentation.
The only variable element is color. The specimens
from the southwestern Indian Ocean are rather colored
with purple or pink (like F assimile). In the
westernmost Pacific, the colors are rather brown, and in
Japan they are entirely vellow to light orange,
sometimes almost white. Two specimens from
Singapore, in AMS, are entirely white.
Differences from F” assimile: Three easily observed
main characters of ! rubicundum. among others.
separate it from }° assimile:
1) Beaded structure of the ribs in the young
specimens; flanks of nbs are smooth in F assimile.
2) Large obliqüe scales on top of ribs in MPQ in
adult specimens. tops smooth in v. assimile (compare
Fig. 25 and Fig. 27).
3) Strongly marked asymmetrical herringbone
patterns on the nbs in the median and anterior parts of
the shells in adult specimens (no herringbone in F°
assimile).
APEX 13(3): 111-125, 20 juil. 1998
121
APEX 13(3} 111-125, 20 juil. 1998
Differences from 1° vertebratum: À beaded double
ornamentation readily comparable to the one in |”
rubicundum also exists in the Australian
vertebratum (Jonas, 1844) [!° reeveanum (Dunker,
I852)], wluch 1s also of a comparable reddish color
However this species belongs to the species-group of
1° flavum (Linné) (see VIDAL 1997b) and significant
characters separate the two shells. F° vertebratum 1s
characterized by
1) Average number of ribs: 29, instead of 36
2) Juvenile ribs in PQ rather symmetrical, without
any anteror sharp edge. but with secondary ornaments
(small scales and ndges, in addition to the main scales,
on the margins of top).
3) Ribs always rather symmetrical.
+) Interstices much more strongly striated.
5) Double beaded ornamentation regular and
constant, not limited to the voung parts of the shell.
6) Herringbone pattern on the nbs rare.
Vasticardium rhegminum
(Oliver & Chesnev, 1997).
Figs 15-17, 29-30
Acrosterigma n. sp. (b) Oliver. 1995: 246. fig. 1092.
Trachycardium (Acrosterigma) rhegminum Oliver &
Chesnev, 1997: 69, Figs 39-42.
Type material.
Five immature bivalved shells. live collected from off
Ras Madrakah. Gulf of Masirah. Oman. coll. "John
Murray" Expedition Sta. 53, 19°22'36"N-57°'53'00"E.
13.5 m. Holotype in BMNH. Four paratypes. three in
BMNHI, one in NMW reg. NMW.Z.1994
Description.
Shell medium-sized , solid, and rather heavy. nearly
equilateral, and only very slightiv oblique (nbs only
shghtly curved backwards in projection). Outline ovate.
generally elongated (mean L/H= 0.83: range 0.77-0.87)
and moderately tumid (mean W/L= 0.84: range 0.80-
0.90). Anterior dorsal slope longer and stecper than
posterior, and posterior margin slightlv straightencd.
Lunular area small, elongate and purple colored.
External color white to beige with an irregular
pattern of pink (gencrally dominant) to light purple-
brown. internally, margin tunged orange-pink along
posterior edge and lightly suffused pink or orange-pink
over umbonal cavity. with two umbonal rays.
122
Revision of the l'asticardium assimile species group
VIDAL
Hinge wide and strong in adult shells, markedly
acutely angled compared to other species of group (< A
range 105-115°) and of different asymmetry (ratio D
about 0.95). Foundation of anterior laterals not hook-
shaped, and à long, elevated medial nb (umbonal
support) occurs in umbonal cavity of all shells.
Mean rib number 40.2, range 38-43.
Rib morphology:
In juvenile PQ, initial rib morphology conforms that
of other species of this group: nbs asymmetrical, with
anterior edge overhanging interstice, and a sharp
continuous margin Jutted in front of scales: nb tops
bear large somewhat tubercular, slightly twisted scales
obliquely set; interstices are as wide as ribs. In adult
PQ. scales flatten with growth, become round and
widen. occupying all the top area, and forming
successive oblique "waves", touching one another:
these waves form crenulations on posterior margin of
nbs, but on anterior margin they gradually disappear,
forming a straight line which progressively overhangs
interstice above original nb margin (Fig. 29). This
results. on anterior margin, in à double overhanging
edge divided by a well delineated thin furrow (Fig. 30).
In juvenile median part of shell, mgh ribs with
roundily triangular top zones and crenulated margins
overhang smooth and flat-bottomed interstices: flanks
smooth. not separated from interstices. In adults (Fig.
29). ribs become very high. and progressively
asymmetrical, with vertical smooth flanks, and smooth
flat tops with more or less regularly crenulated margins
overhanging interstices. Interstices deep, with flat.
smooth bottoms, as wide as or wider than ribs.
In AQ, and as early as in anterior part of MAQ in
very adult shells, marginal crenulations join from one
side to other of rib tops, forming transverse oblique
nidges. rarely with a herningbone pattern.
Material examined.
In addition to the type specimens, the following lots:
A) 43 valves. as type series, BMNH.
B) 15 valves, from storm beach opposite the British
Eastern Relay Station camp, Masirah Island, in NMW
coll. Oliver & Chesney Nov 1992 Reg.
NMWZ.1993 XZ.
C) 1 valve from Masirah area. in BMNH coll. Biggs
1967.
D) 3 small valves from Masirah Island, in MNHN coll.
Day 1994
E) + valves from Masirah Island, in MNHN coll. Fuller
1968.
F) 1 bivalved specimen from Masirah arca. in MNHN
leg Suc Hobbs 1995 (Figs 15-17).
G) 1 bivalved specimen from Masirah arca, in Sue
Hobbs’ private coll.
VIDAL Revision of the J'asticardium assimile Species group APEX 1363): 111-125, 20 juil. 1998
Measurements of }° rhegminum
Height Length Width L/H WI/L ratio D | angle A Ribs
(mm) (mm) (mm) "
Holotype 34.7 28.9 24.4 0.84 0.84 38
53.6 39.4 36.4 0.73 0.92 0.94 105 39
38.9 345 0.75 0.89 0.95 105 41
46.9 (42.0) 0.74 0.90 110 42
MNHN (Fuller) 34.6 (32.4) 0.77 0.94 115 39
MNHN (Fuller) 22.6 (20.0) 0.84 0.88 1.00 38
Largest specimen observed, the above right valve from Masirah Island, in MNHN coll. Fuller (H= 63.4 mm).
Distribution.
To date, this species is known only from the Bay of
Khaliy (Masirah), on the southern coast of Oman,
where it is sympatric with l” assimile.
Remarks.
Variability of l. rhegminum: This species 1s
remarquably constant from one specimen to another, in
all growth stages.
Differences from the three other species of the group:
The average rib number of F° rhegminum 1s higher than
in } assimile and l rubicundum (40.2 vs 33.4 and
35.6). This species differs also from the three other
species of the species group by its very much smaller
angle À, its characteristic "wavy" posterior zone with a
furrowed anterior edge, and also by its very high
regular ribs with smooth flanks and wide smooth
interstices in the other parts of the shell.
Differences from some other species of l’asticardium
with high nbs and wide interstices: The very special
character of PQ is sufficient to separate l”! rhegminum
from several other forms which also have very high
ribs and wide interstices, such as some subspecies of F”
elongatum (see VIDAL 1993), F! fidele (see VIDAL
1992), and some subspecies of F. orbita (see VIDAL
1997a).
Vasticardium thomassini sp. nov.
Figs 21-24, 28, 32.
Type material,
Holotype (Figs 21-24, 28, 32), a shell from Tulear area
(Madagascar), Thomassin's sta. D21 , 43°49'E-2393/'S.
45m, in MNAN. Paratype 1, a left valve from the same
station, in MNHN. Paratype 2, a left valve from Tulear
area, Thomassin's sta. 211, 43°37'E-23°31'S, 12m, in
MNEAN. Paratype 3, a shell from south side of Pwakun
Is, W.Zanzibar, ca. 25 m. in ANSP 213809. Paratype 4,
a right valve from between Port Shepstone and Port
Edward, Natal (South Africa), in MNHN, leg Eavranos
ex Polak.
Description.
Shell small to medium, regularly subovoid, almost
ellipsoidal in shape an almost equilateral, except for a
small truncation in PQ: holotype has also a small
truncation in MPQ. Rüibs generally straight, rarely
slightiy curved in projection. Adult shell little
elongated (L/H range in the three largest shells 0.81-
0.83) and relatively tumid (W/L range in the same
shells 0.82-0.87). Lunular area well delineated, rather
narrow, dark purple colored.
Exterior splashed with yellow to orange, with darker
irregular stripes, interior white with an umbonal area
yellow to orange, sometimes with two darker rays:
margin not colored.
Hinge moderately arched (<A circa 135°) and almost
symmetrical (ratio D circa 1.0 or a little more).
Posterior cardinal in left valve (4b) rather long and
high, with a pointed top. Foundation of anterior lateral
in right valve moderately hook-shaped. No medial
ridge in umbonal cavity.
Rib number ranges 42-45.
Rib morphology:
In juvenile PQ, nibs rather low and assymetrical with
anterior edge overhanging interstice with a sharp
continuous margin jutted in front of scales or dividing
into successive transverse bent lamellae; nb tops bear
large, somewhat tubercular, slightly twisted scales,
obliquely set. Except for the two last ones, interstices
are narrow. In adult PQ (Fig. 28), anterior edge of ribs
becomes less sharp and does not overhang interstice:
the previous thin bent lamellae of anterior edge become
small flank scales with a different obliquity from the
one of the main scales, forming an angle with them.
and being about twice more numerous.
In juvenile median part of shell, nbs are, at first,
flatly rounded, smooth, with progressive appearance of
small serrations at their lower part, separated from
interstice which is finely striated or notched. Then
basal serrations enlarge, become overhanging and nbs
progressively become roundly triangular. In adult MPQ
(Fig. 28), ribs remain about same as in PQ with same
assymmetry (anterior flank shorter and steeper), but
posterior flank becomes finely ridged, when anterior
123
APEX 13(3): 111-125, 20 juil. 1998
small scales become more numerous and progressively
change, onwards, into small, tlun flank ridges, the
equivalent of PQ main scales change into tubercles on
top of ribs. In adult middle of shell (anterior part of
MPQ and posterior part of MAQ,. Fig. 32), nbs become
about symmetnçal and triangular, with both flanks
bearing numerous tlun ridges not reaching interstice.
shorter and more numerous in anterior flank, forming
an hernngbone pattern with the posterior ones. Top
zone progressively loses, onwards. its tubercles and
becomes smooth.
Revision of the Fasticardium assimile species group
VIDAL
In AQ. ribs become more asymmetrical (posterior
flank shorter) and shghtly overhanging and lateral
ndges progressively join together to form top
imbniçated rugac.
Material examined and distribution.
The type series, consisting only of 1 valve from S.
Natal, 1 specimen and 2 valves from S.W. Madagascar,
1 specimen from Zanzibar. In these three localities 1t 1s
sympatric with F. assimile and F. rubicundum.
Measurements of 1° thomassini
Height Width
ratio D
(mm) (mm) (mm)
34.0 27.7 235
Paratype2 32%) 26.7 (22.0)
Paratype3 15.4
Paratype4 12.0
23.0 202
18.5 16.1
Bernard A. Thomassin conducted extensive field work
in Tulear area (S.W. coast of Madagascar). between
1963 and 1972, for ecological studies (see THOMASSIN,
1978).
Etymology.
Remarks.
Differences from Ÿ. rubicundum: V. thomassini is very
close to l. rubicundum as far as shape, morphometric
data, lunule, hinge are concerned; it differs from the
specimens from Africa in absence of purple coloring in
the exterior, and is closer, in that domain, to the
yellow-orange forms of Japan (compunctum). I differs
from all the forms of l”! rubicundum in:
(1) Number of ribs ranging 42-45, when the maximum
in F rubicundum 1s 39.
(2) Rib morphology (compare Figs 28 and 32 with Figs
27 and 31): presence in }. fhomassini of secondary
anterior small scales in PQ and much more numerous
lateral serrations or ridges in the rest of the shell: in F°
thomassini the interstices in the median part of the shell
are much narrower (see Fig. 32); in this median part,
the small lateral ridges are much more numerous and
form a better herringbone pattern in l. fhomassini:
absence also of the "double-beading" of the ribs in the
median part of the juvenile shell which characterizes J”
rubicundum.
ACKNOWLEDGEMENTS. I am especially grateful to
Philippe Bouchet and Bernard Métivier, MNHN Paris,
for their help in many ways. I thank the following
people for allowing me to visit Museums, for loans.
124
informations, and donations to the MNHN: Ilan Loch
(AMS, Sydney), Gary Rosenberg (ANSP,
Philadelphia): Robert Cowie (BISHOP, Honolulu);
Kathie Way (BMNH, London), J. van Goethem and
Claude Massin (IRSNB,. Bruxelles); James McLean
and Lindsey Groves (LACM, Los Angeles), Yves Finet
(MNHG, Genève), R. N. Kiülburn (Natal Museum.
Pietermaritzburg), E. Gittenberger and R. Voskuil
(RMNH. Leiden): Graham Oliver (NMW, Cardiff):
Kevin Lamprell and John Stanisic (QM, Brisbane);
Alan Kabat and M. G. Harasewych (USNM,
Washington), Shirley Slack-Smith (WAM, Perth);
Robert Moolenbeek (ZMA. Amsterdam): Tom Schiotte
(ZMUC, Copenhagen). Very special thanks to Mrs Sue
Hobbs of Cape May, New Jersey USA, who frienly
helped me with her large collection and great
knowledge of Cardnds, and made several donations to
MNAN. I am also indebted to Mrs Elizabeth Ruggerni,
for editing and correcting the english of the manuscript.
REFERENCES
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recent shells from Java Zoologische mededelingen
25: 140-154.
BosCH, D. & E. BosCH. 1982. Seashells of Oman.
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BRAGA, J.M. 1952. Materiais para o estudio da fauna
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DALL, W.H. 1900. Synopsis of the Family Cardiidae
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VIDAL
DRIVAS, J. & M. JAY. 1988. Coquillages de la Réunion
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FISCHER-PIETTE, E. 1977. Révision des Cardiidae
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Série, Série À, Zoologie, Tome 101, 212 pp.
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n°1 & 2, 186 pp.
IREDALE, T. 1927. New molluscs from Vanikoro.
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JONAS, JH. 1844. Vorläufige Dianosen neuer
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KEEN, A.M. 1945. List of shells collected in vicinity of
Oro Bay, New Guinea by Lt. Col. Hubert C. Schenk
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Southern California 10(49): 36-38.
KIRA, T. 1955. Coloured illustrations of the shells of
Japan. Hoïkusha, Osaka, 204 pp.
KIRA, T. 1959. Coloured illustrations of the shells of
Japan. Enlarged & Revised Edition. Hoiïkusha,
Osaka, 239 pp.
KIRA, T. 1962. Shells of the Western Pacific in colour.
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LINNAEUS, C. 1758. Systema Naturae. Salvius, Ed. 10,
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MELVILL, J.C. & R. STANDEN. 1899. Report on the
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of Prof. A.C. Haddon to the Torres Straits in 1888-
89. Journal of the Linnean Society of London 27:
150-206.
OLIVER, P.G. 1992. Bivalved seashells of the Red Sea.
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ni
[ee]
WA
HOUART
Trophon iarae n.sp.
APEX 13(3): 127-130, 20 juil. 1998
Description of Trophon iarae n.sp., a new muricid from
southern West Atlantic (Gastropoda: Muricidae)
with illustration of related species
Roland HOUART
Research Associate, Institut royal des Sciences naturelles de Belgique
Département des Invertébrés Récents
Rue Vautier, 29, B-1000 Bruxelles, Belgium
KEY WORDS. Gastropoda, Muricidae, southern West Atlantic, 7rophon n. sp.
ABSTRACT. 7rophon iarae n.sp. is described and compared with 7. plicatus (Lightfoot, 1786), T.
acanthodes Watson, 1883, T. varians (d'Orbigny, 1841), and 7: geversianus (Pallas, 1774).
INTRODUCTION
Some time ago, I received from Pedro H. Kahvedjian, a
shell collected off Uruguay. After comparison with
specimens in my collection, I identified it as a form of
Trophon plicatus (Lightfoot, 1786). Recently, Prof.
Emily Vokes asked 1f I would agree to study, and
eventually to describe a 7rophon sp. from Brazil. The
specimen was sent to her by Prof. E. Rios, but because
of time constraints, she delayed its study. I accepted of
course, always being very excited by new, and unusual
material.
The specimen I received is gorgeous, and at first sight,
I also felt it was undescribed. Moreover, when
comparing it with my reference material, I came back
to the unusual form of 7: plicatus from Uruguay, and
immediately I saw that it was the same species. The
specimen from Uruguay is smaller, somewhat
damaged, and dead taken. However, now, with the help
of the new material, it is possible to separate it
definitively from the other Magellamic species of
Trophon.
Another specimen of 7: iarae n.sp. was illustrated by
CERNOHORSKY (1977: 117, fig. 18), as TZ. plicatus,
from the Falkland Islands.
The radula of the Brazilian specimen was illustrated by
CALVO (1987: 136, fig. 99), and wrongly identified (as
Tinitially did), as 7: plicatus. However, in her table of
dimensions, CALVO (1987: 174), cites that specimen as
C = 50 mm, L = 35 mm (C = comprimento = length; L
= Jlargura = width). This does not agree with the
dimensions of the shell. Nevertheless, it was confirmed
by Prof. Rios that the radula illustrated by Calvo was
extracted by her from that shell, so that I conclude the
dimensions given by Calvo were probably erroneously
taken from another shell.
SYSTEMATIC ACCOUNT
Genus Trophon Montfort, 1810
Type species by original designation: Murex
magellanicus Gmelin, 1791 (= Buccinum geversianum
Pallas, 1774); Recent, Magellanic region.
Trophon iarae n.sp.
Figs 1-5
Type material.
Holotype n° 6731, Museu Oceanogräfico "Prof. Eliézer
de C. Rios", 74.6 X 61 mm, 1 paratype coll. R. Houart,
53.8 X 39.4 mm.
Type locality.
Off Albardäo, RS, Brazil, collected by fishing boat
"Pescal 2", C. Pontes, 1961. in 55 m.
Other localities.
Off Uruguay, by fishermen (paratype R. Houart); East
of Lively Id, E Falkland Islands, in 79 m
(CERNOHORSKY, 1979).
Distribution.
From the Falklands Islands to Albardäo. RS, Brazil,
living in 55-79 m.
Description.
Shell large sized, up to 74.6 mm in length at maturity
(holotype), heavy, strongly lamellate. Spire moderately
127
APEX 13(3} 127-130, 20 juul. 1998
high, up to 6, broad, strongly shouldered, teleoconch
whorls. Suture adpressed. Protoconch unknown
(broken).
Axial sculpture of teleoconch whorls consisting of
strongly raised, thin lamellae, more strongly developed
on shoulder, producing long, narrow, spinelike
projections. First whorl with 8 lamellae, second to
fourth with 8 or 9, penultimate with 10, last whorl with
13 lamellae. Other axial sculpture of numerous, fine,
narrow, growth striae. Spiral sculpture consisting of a
single caninal cord, forming long, spinelike expansions
at intersection with axial lamellae.
Aperture moderately large, broad, roundly-ovate.
Columellar lip smooth, lip partially erect, adherent at
adapical extremity. Outer lip weakly erect, smooth.
Siphonal canal moderately short, narrow, straight,
open.
Shell entirely white.
Operculum dark brown, roundly-ovate, with terminal
nucleus in lower night.
Radula (in CALVO, 1987: fig. 99): Rachidian with long
central cusp; lateral cusps long with small, inner lateral
denticle; marginal area with several small denticles or
plicae; marginal cusps short.
Discussion.
From 7rophon plicatus (Lightfoot, 1786) (Figs 8-10),
T. iarae n.sp. differs in having a higher spire, and in
having long, almost horizontal, narrow, carinal, open
spines, while the axial lamellae in 7. plicatus end as
short to very short, mostly adapically bent, broad, open
expansions. The shoulder (or sutural ramp) is broad,
weakly convex, and abapically sloped in 7. iarae, but
almost horizontal and narrow in 7: plicatus. The
siphonal canal is narrower and longer relative to the
shell length in 7: iarae. Trophon acanthodes Watson,
1883 (Figs 6-7) also has a broad, weakly convex,
sloped sutural ramp, and narrow, open, carinal spines
as in 7. iarae, but T. iarae differs in other aspects. The
siphonal canal of 7: iarae is twice as short, for a shell
with a same number and same length of spire whorls;
the axial lamellae are more obvious and broader
compared to those of 7: acanthodes which are low, or
almost obsolete; the spiral sculpture is absent in 7.
iarae, while consisting of narrow, conspicuous,
rounded cords in 7. acanthodes.
The two other Magellanic species, namely 7. varians
(d'Orbigny, 1841) (Fig. 12), and 7. geversianus (Pallas,
Trophon iarae n.sp
HOUART
1774) (Fig. 11) differ in many aspects: 7. varians has à
rounded shell, smooth, almost smooth, or ornamented
with coarsely woven spiral cords, without lamellae. 7.
geversianus has more or less, strong, obvious, spiral
sculpture, and usually more numerous, lower, axial
lamellae. 7. geversianus also have a broader aperture.
Both species have a shorter, broader, siphonal canal.
The operculum of 7. iarae nsp., T. plicatus, T.
geversianus, and 7. varians all have a similar
morphology. The radula is also almost identical in 7.
iarae, T. geversianus, and T7. varians, however, it 1s
quite different in 7. plicatus, in having broad, serrate,
lateral cusps (POWELL, 1951; VOKES, 1992).
Trophon acanthodes 1s also different in having a
narrower, triangular operculum with terminal nucleus,
and a broader radula with a shorter, narrower, and more
prominent central tooth.
Etymology.
Named after lara Swoboda Calvo, oceanographer in
Museu Oceanogräfico "Prof. Eliézer de C. Rios", who
extracted and illustrated the radula of the holotype.
ACKNOWLEDGEMENTS. I am very grateful to Prof. E.
de C. Rios, Muséu Oceanogräfico "Prof. Eliézer de C.
Rios", Rio Grande. RS, Brazil, and to Prof. EH.
Vokes, Tulane University (retired), for giving me the
opportunity to study that specimen, and to Mr. P. H.
Kahvedjian (Montevideo, Uruguay) for the other
specimen, sent many years ago. Prof. E.H. Vokes, and
an anonymous reviewer also add some useful remarks
on the manuscript.
REFERENCES
CaALvo, LS. 1987. Räadulas de Gastropodes Marinhos
Brasileiros. Editora da furg: 1-201.
CERNOHORSKY, W.O. 1977. The taxonomy of some
southern ocean Mollusca (Gastropoda) mainly
Antarctic and Subantarctic. Rec. Auckland Mus. 14:
105-119.
POWELL, A.W.B. 1951. Antarctic and Subantarctic
Mollusca: Pelecypoda and Gastropoda. Discovery
Reports 26: 47-196.
VOKES, EH. 1992. Argentine trophons revisited - or
Dr. Powell, I owe you an apology. Amer. Conch.
20(2): 3-4.
Figs 1-3. Trophon iarae n.sp. Off Albardäo, RS, Brazil, 55 m, Museu Oceanogréfico "Prof. Eliézer de C. Rios", 74.6
x 61 mm (whitened, photographs E. VokEs), holotype n° 6731 (siphonal canal damaged after the photograph was
taken). Fig. 4. Distribution of Trophon jarae n.sp.
128
HOUART Trophon iarae n.sp. APEX 13(3): 127-130, 20 juil. 1998
129
APEX 13(3): 127-130, 20 juil. 1998 Trophon iarae n.sp HOUART
—
Fig. 5. T. jarae n.sp. Off Uruguay, 54 mm, paratype coll. R. Houart Figs 6-7. T acanthodes Watson, 1883. Off
Argentina, 59° S, 38° W, 86 mm, coll. R. Houart. Figs 8-9. T. plicatus (Lightfoot, 1786). Rocha, Uruguay, 63.4 mm,
coll. R. Houart. Fig. 10. 7. plicatus (Lightfoot, 1786). Rocha, Uruguay, 344 mm, coll. R. Houart. Fig. 11. T.
geversianus (Pallas, 1774). Straits of Magellan, 93 mm, IRSNB 1G 10591. Fig. 12. 7. varians (d'Orbigny, 1841).
Patagonia, 76 mm, syntype MNAN.
130
KANTOR & TURSCH
Oliva ouini
APEX 13(3): 131-153, 20 juil. 1998
Oliva ouini, a new species from Hansa Bay, Papua New Guinea,
with notes on the anatomy of O. oliva (L., 1758) !
Yuri KANTOR
A. N. Severtzov Institute of Problems of Evolution,
Lenin Avenue 33, Moscow 117071, Russia.
and
Bernard TURSCH
Laboratoire de Bio-Ecologie, Faculté des Sciences, Université Libre de Bruxelles,
50 av. FD. Roosevelt, 1050 Brussels, Belgium.
KEY WORDS. Olividae, Oliva ouini, sp.nov., Oliva oliva, anatomy, shell morphometry.
ABSTRACT. Oliva ouini, sp. nov. from the Western Pacific Ocean is described and compared to its
closest relative: ©. oliva (L., 1758). The shells of the two species differ by several characters and are
completely separable in scatter diagrams. The anatomy of the two species is described and shown to
differ for several features.
RESUME. Ofiva ouini, sp. nov. de l'Ouest de l'Océan Pacifique est décrite et comparée à l'espèce la
plus proche: ©. oliva (L., 1758). Les coquilles des deux espèces diffèrent par plusieurs caractères et
sont complètement séparables en diagrammes bivariés. L' anatomie des deux espèces est décrite; elle
diffère par plusieurs points.
INTRODUCTION.
Since 1973, the various biotopes of Hansa Bay (Papua
New Guinea) have been systematically and repeatedly
explored for their Oliva fauna. The first specimens of
the small species described below were found in May
1992, by SCUBA diving in depths of 6-7 m, around the
bow of a small World War II Japanese wreck locally
known as the "Small Awar wreck". When sifting the
sediment with a small hand dredge (mesh: 8 mm), Mr.
Jean-Marc OUIN (then Manager of Laing Island
Biological Station) noticed that some small, dark Oliva
were escaping through the mesh. These specimens
were unusually fast and agile, re-burying very rapidlv
in the sediment, in which -being highly cryptic- they
vanished if not caught immediately. Albeit quite
elusive, the species is not rare around the wreck and,
during subsequent years, a total of over 30 specimens
has been observed.
The discoverer immediately suggested it was a new
species. One of us (BT) was long hesitant because the
shells somewhat resemble juveniles of the highly
variable Oliva oliva (L., 1758), especially the
Melanesian variety /ongispira Bridgman, 1906, also
present in Hansa Bay. The new form was recognizable
at first sight, but so are many local forms of ©. oliva, a
species known to exhibit extreme inter-population
differences even within short distances (see TURSCH
1994). The protoconchs are quite similar. The
possibility of dealing with an unusual, isolated
population of ©. oliva was increased by the fact that all
specimens known at the time were not syntopic with O.
oliva and came from an area of less than 200 square
meters (this objection can now be discarded: the
species has recently been found in Vanuatu). So the
status of the new species remained long undecided [it
was reported as "species ZHB"' in a study of the Oliva
of Hansa Bay (VAN OSSELAER ef al. 1993)].
Although ©. ouini is easily separated from ©. oliva
(L., 1758) by morphometric analysis of the shell (see
below) and lives in another habitat (at least in Hansa
Bay), it was felt that study of the soft parts could
provide independent evidence of distinct specific
status. This would also provide an opportunity of
describing the hitherto unknown anatomy of Oliva
oliva (L., 1758), the type species of the genus Oliva.
The anatomy of Oliva species has indeed been quite
neglected so far, excepted for the works of KÜTTLER
(1913), MARCUS & MARCUS (1959) and KANTOR
(1991).
Family OLIVIDAE Latreille, 1825
Subfamily OLIVINAE Latreille, 1825
Genus Oliva Bruguière, 1789.
Oliva oliva (L., 1758).
SHELL.
The shell of ©. oliva is extremely variable, with the
consequence that at least two different species form an
"O. oliva complex" and are usually confused by
authors. Their distinction by morphometric analysis
and their geographical distribution have been treated in
! This is paper 30 in the series Studies on Olividae and Contribution n° 350 from Laing Island Biological Station.
131
APEX 13(3}: 131-153, 20 juil. 1998
TURSCH, MISSA & BOUILLON (1992). O. oliva presents
a very large inter-population variation: many
populations (even within very short distances) can be
easily separated, although the whole set of populations
forms one unbroken morphological continuum (see
TURSCH 1994). The variability of ©. oliva is further
increased by non isometric growth: the shape of the
shell varies with age (see TURSCH 1997). The
distribution of shell sizes within one population is very
uneven and has been shown to remain constant
throughout the year (see TURSCH, OUIN & BOUILLON
1995).
ANATOMY.
17 specimens within the shell length range (H) 11.6-
33.4 mm were examined (Table 3). The radulae of 5
specimens were studied by scanning electron
microscopy (SEM).
External anatomy. The body of a specimen with H:
15.8 mm consists of 3.75 postnuclear whorls (PI. 3,
Figs. A-B), the mantle cavity spanning ca. 1/3 whorl.
In alcohol-preserved specimens, the body is pale
yellowish, unpigmented. The foot is thin, folding
longitudinally during fixation, posteriorly it forms a
pouch (PI. 3, Fig. A - fp). The length of the columellar
muscle varies from 1 whorl (specimen H: 15.8 mm) to
1.5 whorls (specimen H: 33.4 mm).
Mantle cavity. Mantle edge even. Mantle rather thick,
although the osphradium and the ctenidium are seen
through it. Siphon long with smooth edges, extending
substantially [33% to 44% L (lip length)] beyond the
mantle edge.
Osphradium yellowish, bipectinate, becoming
relatively smaller as the animal grows (compare PI. 3,
Fig. D and PI. 4, Fig. D). It varies from 83% (specimen
H: 13.1 mm) to 22% (specimen H: 33.4 mm) of the
width and from 84% (specimen H: 12.3 mm) to 62%
(specimen H: 33.4 mm) of the length of the large,
deeply hanging ctenidium. Osphradium asymmetrical:
there are more lamellae on its right side than on the left
(Table 3); the total number of lamellae increases as the
animal grows. The ctenidium occupies nearly 4/5 of the
mantle length. The ctenidium becomes wider and the
lamellae become relatively fewer as the molluscs
grows (compare PI. 4, Figs. E and F). Hypobranchial
gland moderately glandular, forming very low
transverse folds. Anterior mantle tentacle flat, usually
much shorter than the siphon. Posterior mantle tentacle
not pigmented, short and measuring 15% to 25% of H.
Mantle lobe small, concave.
Digestive system. One specimen (male, H 15.8 mm)
was preserved with its proboscis everted (PL 3, Fig. E).
The proboscis is not long (11% of H when contracted
to 28% of H when extended), narrow (length/diameter
— 4,3-8) and lies within the thin-walled proboscis
sheath. The proboscis can be highly retracted during
invertion, so that the buccal mass (PI. 5, Fig. E - od)
and the radular sac (rs) protrude beyond its posterior
end.
Several thin retractor muscles are attached to the
middle part of the rhynchodaem (wall of the proboscis
132
Oliva ouini
KANTOR & TURSCH
sheath) when the proboscis is retracted (PI. 5, Fig. E -
pr). During evertion of the proboscis, the entire length
of the rhynchodaem becomes the proboscis walls and
the point of attachment of the retractors shifts inside
the proboscis.
After the proboscis, the oesophagus is rather narrow
and forms a long loop when the proboscis is inverted.
Duning evertion, the loop is completely straightened.
Valve of Leiblein small and poorly delimited from
the oesophagus (PI. 3, Fig. E - vL; PL 5, Fig. E), which
becomes very narrow to pass through the nerve ring.
The nerve ring is large, massive and without visible
borders of ganglia (PI. 3, Fig. E - nr, PI. 5, Fig. E).
After the opening of the duct of the gland of
Leiblein, the posterior oesophagus widens markedly
towards the stomach (PI. 3, Fig. E; PL 5, Fig. E - poe).
Gland of Leiblein medium-sized, tubular, coiled,
very light-brownish, opens into the oesophagus by a
constricted duct which is close to the nerve ring (PI. 5,
Fig. E - dgL) or lies separately on the right side of the
foregut (PI. 3, Fig. E). Salivary glands medium-sized,
ramified-tubular, rounded or elongated. Salivary ducts
rather thick; shortly after leaving the glands (anteriorly
to the valve of Leiblein) they enter the oesophagus
walls and pass inside them.
The unpaired small accessory salivary gland is
partially embedded in the right salivary gland (PI. 3,
Fig. E; PI 5, Fig. E - asg), with a thick duct (dasg)
which passes at the right side of the oesophagus.
The radula consists of 132 (specimen H: 12.3 mm)
to 184 (specimen H: 23.0 mm) rows of teeth, of which
25-42 rows are not yet completely chitinized. Radula
width varies from 0.73% to 1.30% of H (mean: 0.93%;
©: 0.21; n=9). The lateral teeth are of complex shape,
typical for the genus Oliva: subtriangular, slightly
concave plates with narrow base and curved hook-like
tips. The basal part of the rachidian teeth has distinct
borders; in dorsal view the anterior (directed towards
the mouth) edge is clearly convex, semi-elliptical. The
rachidian tooth has 3 cusps, the central one being the
smallest. In young specimens the cusps are very close
to each other (PI. 8, Figs. 1-3); in larger specimens
their spacing slightly increases (PI. 8, Fig. 4).
The radulae of specimens from two populations in
Hansa Bay have been compared: Boro Beach (steep
white beach, strong wave action) and Sisimangum
Beach (gently sloping black beach, moderate wave
action). The two populations are separated by less than
one kilometer but their shells are very different: on
Sisimangum Beach most specimens are very dark, with
short spires while on Boro Beach all specimens are
whitish with long spires (see TURSCH 1994). The
studied specimens of the Boro Beach population (PI. 8,
Figs. 5-8) appear at first glance to have broader
rachidian teeth. This illusion stems from a change in
general tooth shape (it is relatively shorter) but the ratio
of width to H of the rachidian teeth is practically the
same as in the specimens of the Sisimangum
population. In the Boro Beach specimens this ratio is 0.
27-0. 33%, while in Sisimangum specimens it is 0.23-
KANTOR & TURSCH
0.31%. The rachidian teeth of the Boro Beach
specimens have more widely spaced cusps and present
minute serrations between the cusps (PI. 8, Figs. 7-8).
For the moment we do not know if this is an
ontogenetic change because we have no young
specimens from Boro Beach (where collecting
conditions are rough).
Stomach small, its size and shape differ greatly
among individuals and probably depend on
physiological conditions. The stomach has a rather long
caecum and a digestive gland with a single duct, which
opens just at the entrance of the oesophagus (PI. 4, Fig.
C; PI. 5, Fig. F) (the entrance of the oesophagus is not
clearly seen on PI. 4, Fig. C). The stomach has a small
posterior sorting area and well pronounced typhlosoles.
Anterior sorting area not defined.
Rectal gland absent.
Reproductive system. The gonad, together with the
digestive gland, occupies the upper whorls of the
visceral mass, starting at the level of the nephridium.
The gonad is usually overlaid by the digestive gland
and is sometimes not seen from the outside (PI. 5, Figs.
À, D). Penis in mature males is large, simple,
terminating in more or less long prong (PI. 5, Fig. G)
which is absent in immature males (PI 5, Fig. H).
Accelerated growth of the penis probably occurs at a
shell length of about 14 mm. À male with H: 13.1 mm
still had à penis 0.08 mm long (penis length 0.6% H),
while male with H: 14.1 mm already had a penis 6.25
mm long (penis length 44% of H), although not fully
formed (PI. 5, Fig. H). Afterwards, the relative length
of the penis remains more or less the same throughout
life, or even can become relatively shorter (penis length
34% of H in specimen with H: 25.8 mm; 41% of H in
specimen with H: 27.0 mm). The shape changes
(appearance of the prong) and the seminal duct become
well distinct and visible through the penis walls (PI. 5,
Fig. G). Some males with H: 16.0 mm may still remain
immature.
The maturation of the females occurs probably at
the same shell size. The smallest mature female seen by
us was H: 16.2 mm. It can thus be concluded that ©.
oliva reaches sexual maturity at a shell length of at
least 15 mm.
Ontogenetic changes. Besides sexual maturation, the
only significant ontogenetic change noticed by us is the
relative size of the osphradium and the ctenidium. In
young specimens the osphradium has nearly the same
size as the ctenidium; it becomes much smaller in
grown-up specimens.
Oliva ouini sp. nov.
"Oliva sp. ZHB"; Van Osselaer & al. 1994: 30.
Type Material.
Holotype (H: 15.62 mm; D: 5.91 mm): Natural History
Museum, London [BM(NH)] (PI. 1, fig. 1).
Paratype 1 (H: 13.50 mm; D: 5.51 mm): Institut Royal
des Sciences Naturelles de Belgique, Brussels
(LR.N.S.B.) (PL 1, Fig. 2).
Oliva ouini
APEX 13(3): 131-153, 20 juil. 1998
Paratype 2 (H: 13.59 mm; D: 5.57 mm): Zoological
Museum, Moscow State University (ZMM) n°.Lc
23326 (PI. 1, fig. 3).
Paratype 3 (H: 13.66 mm, D: 5.64 mm): United States
National Museum, Smithsonian Institution (USNM)
(PL. 1, fig. 4).
Paratype 4 (H: 12.91 mm; D: 5.04 mm): Muséum
National d'Histoire Naturelle, Paris (MNHN) (PI. 1,
fig. 5).
Paratype 5 (H: 12.05 mm; D: 4.97 mm): Australian
Museum, Sydney (PI. 1, Fig. 6).
Type locality. Hansa Bay, Papua New Guinea
(4°10'30" S - 144°52'47" E). Near bow of "Small Awar
Wreck", grey sand, 6-7 m.
SHELL.
SIZE: up to about 17 mm.
GENERAL SHAPE: biconical, elongated.
SPIRE: conical, elevated.
PROTOCONCH: greyish-white to dark purplish-grey.
Nuclear whorls: mean 3.6. Lower part of last nuclear
whorl purple. Transition to teleoconch straight, well
defined.
SPIRE WHORLS: profile flat. First postnuclear whorl
white. Subsequent whorls with long axial stripes.
FILAMENT CHANNEL: rather narrow, deep, olack.
BODY WHORL:
SHELL BACKGROUND: yellowish-cream to whitish.
COLOUR PATTERN: Fine to very fine pattern of brown
zigzags, in many cases coalescing into nearly solid
brown zones. The shell background is then seen
mostly in triangular zones, often coalescing into axial
series and commonly delineated with a darker brown
line. The zigzag pattern is reinforced by darker
chevrons, frequently arranged into axial series. In
addition, most specimens have bold dark brown axial
lines, following growth lines.
SUBCHANNEL PATTERN: long, dark radial strokes,
fainting adapically and often coalescing into a
continuous line.
COLUMELLA: somewhat translucent, light purplish grey
to flesh, white in faded specimens. 8 to 9 (generally
8) very strong oblique, parallel columellar plications
forming regular, rounded columellar teeth.
FASCIOLE: whitish-grey with dark-brown spot at tip.
SUPRAFASCIOLAR BAND: dark grey, patterned with
coarse, curved, dark lines in its lower zone,
sometimes extending in the upper zone.
APERTURE: chocolate-brown. Inner margin of lip
darker. Edge of lip beige. External contour of lip
slightly angulate (bulging) near adapical third of
aperture. The elongated aperture is rather distant from
the filament channel of the previous whorl.
Quantitative data. The meaning of the measurements
used in this work is sketched in Figs. 1 and 2. The
linear teleoconch measurements H, L, D and LW, the
number of nuclear volutions NW and the number of
post-nuclear volutions PNW were defined in TURSCH
& GERMAIN (1985); the linear protoconch
measurement RESS was defined in TURSCH &
133
APEX 13(3): 131-153, 20 juil. 1998 Oliva ouini
GERMAIN (1986), the linear protoconch measurement
PAT17 was defined in TURSCH & GERMAIN (1987).
Some operational quantitative charactenstics are
given in Table 1. As usual in species belonging to the
genus Oliva (and contrary to widespread assumption),
the most variable character is the relative height of the
spire (H-L)/H.
Morphometric separation from ©. oliva. This was
effected by comparing 12 specimens of ©. ouini (11
with intact protoconch) from the type locality to 100
specimens of ©. oliva (L., 1758). These 100 specimens
include 10 specimens of each of 10 different local
populations, covering much of the inter-population
variability of the species. At least half of each local
sample consisted of shells less then 25 mm in length, to
avoid the risk of size-related, artificial separations. The
local phena, defined in TURSCH, MIssA & BOUILLON
(1992), are: phenon 74 (W. Thailand), phenon 40
Protoconch (nr = 11)
NW
Teleoconch (7 = 12)
D/H
D/L
0.67 _| 0.030
Ds enesess co en 0S ENS ROM UT EE
EE 2 EN En EU 0.024
0.87
KANTOR & TURSCH
(W. Australia), phenon BA (Indonesia, Bali), phenon
SR (Sn Lanka), phenon //B (Papua New Guinea, Hansa
Bay), phenon W/B (Indonesia, West Java), phenon
WJA (Indonesia, West Java), phenon S/ (Indonesia,
South Java), phenon MB (Papua New Guinea, Milne
Bay) and phenon PA (Philippines).
O. ouini can be completely separated from the
morphological continuum of ©. oliva (L., 1758). Only
two examples will be given here: the scatter diagram of
L/LW vs. D/H (see Fig. 1) and the scatter diagram of
L/PNW vs.PATI7/RESS (see Fig. 2).
The wide dispersion observed for the larger values
of L/PNW in Fig. 2 corresponds to the non-isometric
growth pattern of ©. oliva, causing a marked increase
of the relative length of the lip in large specimens (see
TURSCH 1997).
3.90 0.180
23 %
47%
43%
3.99%
3.3 %
17%
Table 1. Some morphometric characteristics of O. ouini sp. nov. All specimens from type
locality. S.D.: standard deviation; C.V. : coefficient of Variation (100*SD/mean).
Diagnosis. À few easy identification tips are sketched
in PI. 2, Figs. 1,2.
Discussion. At first glance, ©. ouini is similar in shape
to some populations of ©. oliva. The most obvious
difference is the presence in ©. ouini of 8-9 very
strong, oblique columellar plications (see PI. 1, Figs. 1-
6), which are never observed in ©. oliva (L., 1758) (see
PI. 1, Figs. 10-12) or in the closely related ©. tigridella
Duclos, 1835 (see PI. 1, Fig. 9). The protoconchs of the
two species are very similar (see PI. 2, Figs. 3, 6) but
the ratio PATI7/RESS is mostly smaller in ©. ouini
(see Fig. 2). For the same size of shell lip, ©. ouini has
generally more postnuclear whorls than ©. oliva
(smaller L/PNW). The external contour of the lip is
slightly more angulate.
O. ouini is immediately distinguished by its
elongated, biconic body whorl from juveniles of the
syntopic species ©. caerulea (Rôüding, 1798) (see PI. 1,
Fig. 7) which has a completely different protoconch
(see PL. 2, Fig. 4) and from ©. concinna Marrat, 1870
which has a somewhat similar protoconch (see PI. 2,
134
Fig. 5) but differs by many morphometric discriminants
(measurements taken on juveniles with H < 25mm,
n=9), amongst others L/LW (mean 0.93, max. 0.94;
min. 0.92; S.D. 0.006; C.V. 0.63 %; compare with
Table 1).
ANATOMY.
Four specimens were examined (Table 1). The radulae
of two specimens were studied by SEM.
External anatomy. The body of a specimen with H:
11.6 mm consists of 2.5 postnuclear whorls, the mantle
cavity spanning ca. 2/3 whorl (PL. 6, Figs. A-B; PL 7,
Figs. A-C). The live animal of ©. ouini is yellow-beige,
with very contrasting dark brown maculations on all
the foot (see PI. 2, Figs. 7-9). In alcohol-preserved
specimens, the body is pale yellowish, siphon, head
tentacles and propodium partially speckled with
brownish spots. The foot is thin, folding longitudinally
during fixation; posteriorly it forms a pouch (PI. 6,
Figs. À, B; PI 7, Fig. B, C - fp). The length of the
columellar muscle is about one whorl.
KANTOR & TURSCH Oliva ouini
> 0.49
H 0.47
0.45
0.43
0.41
0.39
ApFpOOd=6n00O
APEX 13(3): 131-153, 20 juil. 1998
©. ouini
. OliVa :
Phenon THA
Phenon AO
Phenon BA
Phenon SR
Phenon HB
Phenon WJB
Phenon WJA
Phenon SJ
Phenon MB
Phenon PA
Fig. 1. Morphometric separation of ©. ouini sp. nov. from ©. oliva (L., 1758). Scatter diagram of L/LW vs. D/H.
Minimum convex polygons. See text.
PAT17 PAT17 IRESS
D ARESENS
0.52
0.48
0.42
©. ouini
©. oliva :
App ONE & DO O ©
Phenon THA
Phenon AO
Phenon BA
Phenon SR
Phenon HB
Phenon WJB
Phenon WJA
Phénon SJ
Phenon MB
Phenon PA
< PNW
y
Fig. 2. Morphometric separation of ©. ouini sp. nov. from ©. oliva (L., 1758). Scatter diagram of L/PNW
vsS.PAT17/RES5. Minimum convex polygons. See text.
135
APEx 13(3) 131-153, 20 juil. 1998
Oliva ouini
KANTOR & TURSCH
Mantle cavity (PL 6, Fig. D, PI 7, Fig. E). Mantle
edge even. Mantle rather thick, although the
osphradium and the ctenidium are seen through it.
Siphon long with smooth edges, extending
substantially (32% to 40% of L) beyond the mantle
edge.
Osphradium vyellowish, bipectinate, 55% t0110%
(mean: 88%, ©: 0.23) of the width and 74% to 88%
(mean: 81%; ©: 0.07) of the length of the large, deeply
hanging ctenidium. Osphradium asymmetncal: there
radular rows
E radular width
[ee]
shell length, mm
number of radular
number of forming
rachidian width
number of ctenidium
lamellae
E
E
&
0
=
7
ë
3
=
o
e
Los)
©
en
+
ee
ste ler RLE
DM ERE dE
= & = E is =
0
= = © oO = c 0 =
S |S |s 55 | SUIS E
= a © & = TD E
= o ES Le LE ee
ë AOUE E Se
= = 3 & o
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are more lamellae on the right side than on the left
(Table 2) (in average 54 vs. 47). The ctenidium
occupies nearly 4/5 of mantle length. Hypobranchial
gland moderately glandular, forming very low
transverse folds. Anterior mantle tentacle flat, may be
nearly as long as the siphon. Posterior mantle tentacle
not pigmented, moderately long, about half of the lip
length (L). Mantle lobe small, concave.
Rectal gland absent.
ctenidium width, mm
ctenidium length, mm
number of osphradium
lamellae, left part
number of osphradium
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Table 3. Ofiva oliva (L., 1758). Summary of some anatomical characters examined. Papua New Guinea, Hansa
Bay: last two specimens from Boro Beach, all others from Sisimangum Beach.
136
KANTOR & TURSCH
Oliva ouini
APEX 13(3): 131-153, 20 juil. 1998
Digestive system. One specimen (female, H: 13.9 mm)
was preserved with its proboscis everted (PI. 7, Fig. H).
Proboscis not long [17% of H when contracted (PI. 6,
Fig. F) to 25% of H when everted], narrow
(length/diameter — 4.5-7), and lies within the thin-
walled proboscis sheath.
The foregut of ©. ouini 1s generally very similar to
that of O. oliva.
The radula consists of 80 (specimen H: 11.3 mm) to
91 (specimen H: 13.9 mm) rows of teeth, of which 12-
25 rows are not yet completely chitinized. Radula
width varies from 0.91% to1.06% of H (mean: 0.97%;
G: 0.07; n=4). The lateral teeth are very similar in
shape to that of O.oliva. Rachidian teeth: the lateral
sides of the basal part fuse with the subradular
membrane and are inconspicuous. In dorsal view, the
anterior edges of the basal parts are nearly straight or
very slightly concave in the middle. The rachidian teeth
have 3 cusps, the central one being the smallest.
Stomach small, with long caecum and single duct
of the digestive gland. The shape of stomach differs
greatly amongst specimens. We were unable to
examine the stomach anatomy.
Reproductive system. The gonad, together with the
digestive gland, occupies the upper whorls of the
visceral mass, starting at the level of the stomach (PL 6,
Figs. C, H; PL 7, Fig. F - gon). The gonad is not
overlaid by the digestive gland. The penis is large,
bilobed, with a somewhat flattened basal lobe and a
rounded upper lobe which terminates in small curved
seminal papilla (PI. 6, Figs. E, G). The smallest studied
specimen (H: 11.6 mm) had a fully formed penis,
indicating that the specimen was mature. The smallest
studied female (H: 11.3 mm) had large, fully developed
pallial gonoduct (PI. 7, Fig. E - pgon). It can thus be
concluded that ©. ouini reaches sexual maturity at a
shell length of less than 11.3 mm.
Comparison with ©. oliva.
In spite of the general similarity of their anatomy, the
two species differ by at least four independent
anatomical characters.
1. The radula of ©. ouini has a significantly smaller
number of teeth rows (80-91 vs. 132-184 for ©. oliva,
see Tables 2 and 3).
2. The rachidian radular teeth differ in shape: in ©.
oliva the anterior edge is markedly convex (see PI. 8).
in ©. ouini it is nearly straight and even concave in the
middle (see PI. 9).
3. In ©. oliva the gonad starts at the level of the
nephridium (see PI. 5, Fig. B); in ©. ouini it starts at
the level of the stomach (PI. 6, Figs. C,H; PI. 7, Fig. F).
4. The shape of the penis of ©. oliva (see PI. 5, Figs. G,
H) is very different from that of ©. ouini (see PI. 6,
Figs. E°G)!
S. The strong maculations seen on the live animal of ©.
ouini are not observed on the body of ©. oliva which
comes in different colours, generally matching the
ground colour of the shell (see PI. 2, Figs. 10-13).
It was also shown that ©. ouini reaches maturity at
a size at least 3.5 mm smaller than ©. oliva. Other
differences (such as the size of the anterior and
posterior mantle tentacles) are small and may depend
on the conditions of preservation.
OTHER DATA.
Distribution. For years, known only from the type
locality, where it is confined to a very small area. The
distribution range is in fact much wider. Seven nearly
identical specimens originating from Vanuatu {given
locality: "Ambre Isl." (error for Ambrym ?),1-3 m]
have now been identified in the collection of Mr. J.P.
LEFORT (Tahiti) by Dr. Dietmar GREIFENEDER Who
also has one specimen from Vanuatu, "plage" (2.5 m,
black sand).
Habitat. In Papua New Guinea ©. ouini has been
found only in rather calm water, fine grey coral sand,
6-7 m, around one of the ship wrecks in Hansa Bay. It
is there syntopic with other Oliva species, amongst
others ©. caerulea (Rôding, 1798), ©. concinna
Marrat, 1870, ©. reticulata (Rôding, 1798), ©. sericea
(Rôding, 1798). In contrast, ©. oliva is confined to
open sand beaches, exposed to occasional strong surf.
In Vanuatu, ©. ouini is reported from 1 to 3 m
depth.
Etymology. This species is named for our friend Jean-
Marc OUIN, former manager of Laing Island Biological
Station and master in the art of finding elusive Oliva
species.
Acknowledgements.
We are grateful to the Belgian Fonds National de la
Recherche Scientifique (F.N.R.S.) and to BIOTEC.,
S.A. for supporting our research. We thank Dr.
Dietmar GREIFENEDER for much advise.
REFERENCES.
KANTOR, Yu. 1991. On the morphology and
relationships of some oliviform gastropods.
Ruthenica 1(1-2): 17-52.
KÜTTLER, A. 1913. Die Anatomie von 0. peruviana
Lamarck. Zool. Jahrbuch, suppl. 13 (Fauna
Chilensis): 477-544.
MARCUS, E. & E. MARCUS 1959. Studies on Olividae.
Bol. Fac. Filos. Ciencias e Letras Univ. Sao Paulo,
232 (Zool. 22): 96-188.
TURSCH, B. 1994. Studies on Olividae XXI. The scale
of sympatry in the genus Oliva. Apex 9(4): 131-142.
137
APEX 13(3): 131-153, 20 juil. 1998
TURSCH, B. 1997. Non-isometnic growth and problems
of species delimitation in the genus Oliva. Apex 12(2-
3): 93-100.
TURSCH, B. & L. GERMAIN 1985. Studies on Olividae.
IL À morphometric approach to the O/iva problem.
Indo-Malayan Zoology 1: 331-352.
TURSCH, B. & L. GERMAIN 1986. Studies on Olividae.
Il. Further protoconch morphometrical data for Oliva
taxonomy. Apex 1(2): 39-45.
TURSCH, B. & L. GERMAIN 1987. Studies on Olividae.
V. Five additional protoconch characters for Oliva
taxonomy. Apex 2(3/4): 59-68.
Plate 1. Scale bars: 10 mm.
1-6. O. ouini sp. nov. All from type locality.
. Holotype (H: 15.62 mm; D: 5.91 mm): BM(NH).
. Paratype 1 (H: 13.50 mm, D: 5.51 mm): IRNSB.
. Paratype 3 (H: 13.66 mm, D: 5.64 mm): USNM.
. Paratype 4 (H: 12.91 mm; D: 5.04 mm): MNHN.
. Paratype 5 (H: 12.05 mm; D: 4.97 mm): AMS.
© ou p &w ND =
T-
locality.
Oliva ouini
KANTOR & TURSCH
TURSCH, B., O. MissA & J. BOUILLON 1992. Studies on
Olividae XIV. The taxonomic structure of Oliva oliva
(auct.). Apex 7(1): 3-22.
TURSCH, B.. J.M. OUIN & J. BOUILLON 1995. On the
structure of a population of Oliva oliva (L., 1758) in
Papua New Guinea (Studies on Olividae 22). Apex
10(2/3): 29-38.
VAN OSSELAER, C., J. BOUILLON, J.M. OUIN & B.
TURSCH. 1994. Studies on Olividae XVII. The
distribution of Oliva species and the variation of their
colour patterns in Hansa Bay (Papua New Guinea).
Apex 9(2/3): 29-46.
. Paratype 2 (H: 13.59 mm, D: 5.57 mm): ZMM n°. Lc 23326.
. caerulea (Rüding, 1798). Juvenile shell (H: 14.77 mm; D: 6.46 mm), found together with ©. ouini, in type
8. O. concinna Marrat, 1870. Juvenile shell (H: 14.82 mm; D: 5.96 mm), found together with ©. ouini, in type
locality.
9. ©. tigridella Duclos, 1840. Juvenile shell (H: 16.84mm, D: 6.77 mm), Philippines, Cebu.
10-12. ©. ofiva(L., 1758).
10. Juvenile shell (H: 15.81 mm, D: 5.77 mm), Papua New Guinea, Hansa Bay, Sisimangum Beach, low tide.
11. Juvenile shell (H: 19.92 mm; D: 8.33 mm), Vietnam, Nha Trang, Hon Tre Is., low tide.
12. Juvenile shell (H: 15.66 mm, D: 6.43 mm), Sri Lanka, Welligama, low tide.
138
KANTOR & TURSCH Oliva ouini APEX 13(3): 131-153, 20 juil. 1998
holotype paratype l paratype 2 paratype 3 paratype 4 paratype 5
O. ouini sp. nov.
©. caerulea ©. concinna ©. tigridella ©. oliva
139
APEX 13(3): 131-153, 20 juil. 1998 Oliva ouini KANTOR & TURSCH
Plate 2.
1-3. O. ouini sp. nov. All from type locality.
1.-2. Tips for quick shell recognition.
3. Protoconch.
4. O. caerulea (Rôding, 1798). Protoconch. Juvenile shell (H: 14.77 mm, D: 6.46 mm), found together with ©. ouini,
in type locality.
5. ©. concinna Marrat, 1870. Protoconch. Juvenile shell (H: 14.82 mm, D: 5.96 mm), found together with ©. ouini,
in type locality.
6. ©. oliva (L., 1758). Protoconch. Juvenile shell (H: 15.81 mm, D: 5.77 mm), Papua New Guinea, Hansa Bay,
Sisimangum Beach, low tide. Juvenile shell (H: 15.81 mm; D: 5.77 mm), Papua New Guinea, Hansa Bay,
Sisimangum Beach, low tide.
7-9. O. ouini sp. nov. Live animals. All from type locality.
10-13. ©. ofiva (L, 1758). Live animals, different colour forms, all from Papua New Guinea, Hansa Bay,
Sisimangum Beach, low tide.
140
KANTOR & TURSCH Oliva ouini APEX 13(3): 131-153, 20 juil. 1998
aperture distant D)
— from previous
whorl
characteristic
i maculations on
ù lip slightly suprafasciolar band
angled
very strong dark 4
—— columellar ©. ouini ©. ouini
plications
©. ouini
: . Le
©. caerulea ©. concinna ©. oliva
. “à / Fa
©. ouini
©. ouini
©. ouini
Il ©. oliva
O. oliva 13
©. oliva
©. oliva
141
APEX 13(3): 131-153, 20 juil. 1998 Oliva ouini KANTOR & TURSCH
Plate 3. Anatomy of ©. o/iva (L., 1758).
Specimen(o H: 15.8 mm) from Papua New Guinea, Hansa Bay, Sisimangum Beach.
Scale bars: À, B - 2 mm, C-F - 1 mm.
A, B - body removed from the shell.
C - view of visceral mass, showing the shape of stomach.
D - cut-out mantle.
E - anterior part of digestive system from the right side with the proboscis everted, extended.
F - native position of the gland of Leiblein and salivary glands, from the left anterior side.
amt - anterior mantle tentacle nr - nervous ring
ao - anterior aorta os - osphradium
asg - accessory salivary gland par - parapodium
cm - columellar muscle pen - penis
cme - cut mantle edge pmt - posterior mantle tentacle
ct - ctenidium poe - posterior oesophagus
dasg - duct of accessory salivary gland pr - proboscis
ddg - duct of digestive gland prp - propodium
dg - digestive gland re - rectum
dgL - duct of gland of Leiblein s - siphon
fp - pouch of foot sd - salivary duct
gL - gland of Leiblein sg - salivary gland
gon - gonad st - stomach
hg - hypobranchial gland VL - valve of Leiblein
ml - mantle lobe
142
KANTOR & TURSCH Oliva ouini APEX 13(3): 131-153, 20 juil. 1998
Plate 3. Oliva oliva (L., 1758)
143
APE
X 13(3): 131-153, 20 juil. 1998 Oliva ouini KANTOR & TURSCH
Plate 4. Anatomy of ©. o/iva (L., 1758).
A-D -Specimen (Q H: 33.4 mm) from Papua New Guinea, Hansa Bay, Boro Beach.
Scale bars: À, B, D -5 mm,C-2mm,F-1mm,E-0.5 mm.
144
A, B - body removed from the shell.
C - stomach, opened dorsally.
D - cut-out mantle.
E - Shape of ctenidium lamellae. Specimen (H: 13.1 mm) from Papua New Guinea, Hansa Bay, Sisimangum
Beach.
F - Shape of ctenidium lamellae. Specimen (H: 22.0 mm) from Papua New Guinea, Hansa Bay, Sisimangum
Beach.
amt - anterior mantle tentacle oe - oesophagus
c - caecum of the stomach os - osphradium
cm - columellar muscle pgon - pallial gonoduct
cme - cut mantle edge pmt - posterior mantle tentacle
ct - ctenidium prp - propodium
ddg - duct of digestive gland psa - posterior sorting area
dg - digestive gland re - rectum
gon - gonad s - siphon
ht - head tentacles st - stomach
ig - intestinal groove t - typhlosoles
ml - mantle lobe
Oliva ouini APEX 13(3): 131-153, 20 juil. 1998
KANTOR & TURSCH
Plate 4. Ofiva oliva (L., 1758)
145
APEX 1363): 131-153, 20 juil. 1998 Oliva ouini KANTOR & TURSCH
Plate 5. Anatomy of ©. ofiva (L., 1758). Scale bars: A,B - 2 mm, C-F - 1 mm.
Specimens from Papua New Guinea, Hansa Bay, Sisimangum Beach.
A,H-(9 H:14.1 mm) (drawn to the same scale).
B,E-(Q H: 11.43 mm) (drawn to the same scale).
C;,F,G - (0 H: 25.8 mm).
D-(S H:19.2 mm).
Scale bars: 1 mm.
A-D - views of visceral mass, showing variability of the shape of stomach.
E - anterior part of digestive system from the right side, extended.
F - stomach, opened dorsally.
G, H - ontogenetic changes of the penis shape and size.
asg - accessory salivary gland poe - posterior oesophagus
c - caecum of the stomach pr - proboscis
ddg - duct of digestive gland prr - proboscis retractors
dgL - duct of gland of Leiblein psa - posterior sorting area
gL - gland of Leiblein rhd - rhynchodaeum (proboscis sheath)
gon - gonad rs - radular sac
ig - intestinal groove sg - salivary gland
nr - nervous ring t - typhlosoles
od - odontophore vL - valve of Leiblein
ooe - opening of oesophagus into stomach
146
KANTOR & TURSCH Oliva ouini APEX 13(3): 131-153, 20 juil. 1998
Plate 5. Ofiva oliva (L., 1758)
147
APEX 13(3): 131-153, 20 juil. 1998 Oliva ouini KANTOR & TURSCH
Plate 6. Anatomy of males of O/iva ouini sp. nov.
A-F: (0 H: 11.6 mm); G-H: (© H: 11.8 mm).
Scale bars: A,B - 2 mm; others - 1 mm. C, F, H drawn to the same scale.
A,B - body removed from the shell.
C,H - view of visceral mass, showing the shape of stomach.
D - cut-out mantle.
E, G - penis.
F - anterior part of digestive system from the right side, extended. Proboscis inside the rhynchodaeum is
shown by dotted line.
amt - anterior mantle tentacle os - osphradium
asg - accessory salivary gland par - parapodium
cm - columellar muscle per - pericardium
cme - cut mantle edge pmt - posterior mantle tentacle
ct - ctenidium poe - posterior oesophagus
dasg - duct of accessory salivary gland pr - proboscis
dg - digestive gland prp - propodium
dgL - duct of gland of Leiblein prr - proboscis retractors
fp - pouch of foot re - rectum
gL - gland of Leiblein rhd - rhynchodaeum (proboscis sheath)
gon - gonad s - siphon
hg - hypobranchial gland sg - salivary gland
ht - head tentacles st - stomach
ml - mantle lobe vL - valve of Leiblein
nr - nervous ring
oe - oesophagus
148
Oliva ouini APEX 13(3): 131-153, 20 juil. 1998
KANTOR & TURSCH
Plate 6. Oliva ouini sp. nov.
149
APEX 13(3): 131-153, 20 juil. 1998
Oliva ouini
Plate 7. Anatomy of a female of O/iva ouini sp. nov., (H: 13.9 mm).
Scale bars: À, B,C,E-2mm;F,6G,H-1 mm.
A, B - body removed from the shell.
150
C - ventral view of the foot, showing the ventral pedal gland.
D - enlarged dorsal view of the head with proboscis protruded.
E - cut-out mantle.
F - view of visceral mass, showing the shape of stomach.
G - stomach from inner side.
H - anterior part of digestive system, from the right side, extended.
amt - anterior mantle tentacle
asg - accessory salivary gland
cme - cut mantle edge
ct - ctenidium
dasg - duct of accessory salivary gland
ddg - duct of digestive gland
dg - digestive gland
dgL - duct of gland of Leiblein
fp - pouch of foot
gL - gland of Leiblein
gon - gonad
hg - hypobranchial gland
ht - head tentacles
ml - mantle lobe
nr - nervous ring
oe - oesophagus
os - osphradium
par - parapodium
pgon - pallial gonoduct
pmt - posterior mantle tentacle
poe - posterior oesophagus
pr - proboscis
prp - propodium
re - rectum
s - siphon
sg - salivary gland
st - stomach
vL - valve of Leiblein
vpg - ventral pedal gland.
KANTOR & TURSCH
APEX 13(3): 131-153, 20 juil. 1998
Oliva ouini
KANTOR & TURSCH
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Plate 8. Oliva oliva (L., 1758).
1-3: female, Papua New Guinea, Hansa Bay, Sisimangum (H: 11.6 mm).
4 female, Papua New Guinea, Hansa Bay, Sisimangum (H: 20.2 mm).
5-7: male, Papua New Guinea, Hansa Bay, Boro Beach (H: 27.0mm).
8: female, Papua New Guinea, Hansa Bay, Boro Beach (H: 33.4 mm).
152
KANTOR & TURSCH Oliva ouini
APEX 13(3): 131-153, 20 juil. 1998
Plate 9. Oliva ouini sp. nov. All from type locality.
1-3: female (H: 11.3 mm).
4-7: male (H: 11.6 mm).
153
és
LR 4 NC
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VOL. 13 (4)
R. NN. Kilburn
B. Tursch
E. Rolän
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20 DECEMBRE 1998
SOMMAIRE
Description of four new neogastropods of superfamilies
Muricoidea and Conoidea from South Africa
(Gastropoda: Prosobranchia: Neogastropoda)
A simple shell model: applications and implications
A new species of Zebina (Gastropoda: Rissoidae: Rissoininae)
from Yucatan (Mexico)
APEX
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Dr. J. Van Goethem
Prof. G. Vauquelin
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Muséum d'Histoire Naturelle, Genève
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KILBURN
Four new neogastropods
APEx 13(4): 155-160, 20 déc. 1998
Description of four new neogastropods of superfamilies
Muricoidea and Conoidea from South Africa
(Gastropoda: Prosobranchia: Neogastropoda)
R. N. KILBURN
Natal Museum, P/Bag 9070, Pietermaritzburg 3200, South Africa
KEY WORDS. Gastropoda, Columbellidae, Olividae, Drilliidae, Turridae, South Africa.
ABSTRACT. The following four Neogastropoda are described from the continental shelf or slope of
eastern South Africa: Anachis (Suturoglypta) blignautae (family Columbellidae) from off southern
Zululand;, Ancillista depontesi (Olividae), Splendrillia hayesi (Drillüdae), and Turris faleiroi
(Turridae) from the eastern Agulhas Bank. Turris faleiroi is the first temperate-water member of its
genus known.
INTRODUCTION
The discovery of a notable proportion of the more
striking southern African benthic molluscs is due to the
activities of commercial fishing boats. Several such
species, recently submitted to the Natal Museum by Mr
Brian Hayes of Port Elizabeth, prove to be undescribed.
Three of the species described here may be assumed
from their known distribution to be temperate-water
Agulhas Bank endemics, the fourth (an Anachis) may
possibly prove to be a tropical East African element.
Abbreviations
BMNH: The Natural History Museum, London
NMSA: Natal Museum, South Africa
ZMHB: Zoological Museum, Humboldt University,
Berlin.
TAXONOMY
Superfamily MURICOIDEA
Family COLUMBELLIDAE
Genus Anachis H. & A. Adams, 1853
The following species is referred to the subgenus
Suturoglypta Radwin, 1968 [type species (o.d.)
Columbella pretri Duclos, 1846], on account of its
square-cut whorls. Nevertheless, this may prove to be a
homoplasy, as the only three species previously
referred to this subgenus all inhabit the western
Atlantic region. These were discussed by RADWIN
(1978: 340), who elevated Surturoglypta to full genus
status. In the absence of a modern analysis of the
systematics of the Columbellidae, I prefer to follow a
conservative approach.
Anachis (Suturoglypta) blignautae n. sp.
Figs 1-2
Type material.
Holotype NMSA V6140/T1548. Paratype 1, NMSA
V6141/T1549; paratype 2 in B. Hayes collection. All
taken from crayfish traps; paratypes both juvenile.
Type locality.
Off Richards Bay (c. 28°48’S, 32°05°E), 600 m.
Distribution.
Continental slope of southern Zululand, known onl\
from the type locality.
Diagnosis.
Shell fusiform (aperture/total length 0.50), with a
produced, somewhat dorsally bent anterior end, suture
shallow; axial ribs strong, 12-13 per whorl, projecting
most at periphery of last whorl, evanescing on base,
crossed above and below suture by a spiral thread,
which make spire whorls appear almost flat-sided; base
of last whorl with 14 spiral threads; aperture with
smooth margins, columella callus with shghtly raised
outer edge; siphonal canal not indented. Protoconch
papillose, smooth, breadth 0.63-0.70 mm. Pale pinkish-
orange, ribs and subsutural region white, protoconch 1
with a brown spot. Maximum length 11.2 mm.
155
APEX 13(4): 155-160, 20 déc. 1998
Four new neogastropods
KILBURN
Description.
Shell fusiform, of 6 teleoconch whorls, breadtl/length
0.39, with a high, acute, orthoconoïd spire (slightly
cyrtoconoid towards apex) and a tapering, elongated
anterior end, spire whorls rather flat-sided (almost
quadrate), suture shallow. Aperture long (aperture/total
length 0.50), narrow, greatest width at about posterior
third, siphonal canal long and bent dorsally and to left,
base obliquely truncate, termination not indented; inner
and outer lips smooth, inner lip with a moderately thick
callus, whose outer edge is slightly raised on
columella; outer lip sinuous in side-view, evenly
convex medially, shallowly concave below suture.
Sculptured by moderately strong axial ribs, crossed
above and below suture by a spiral lira, base of last
whorl spirally lirate. Axial ribs opisthocline, rather
straight, in cross-section rounded-angular, subequal to
their intervals, suture to suture, evanescing on last
whorl in parietal region, most prominent at periphery
of body whorl; early whorls with 13 ribs, decreasing to
12 on penultimate whorl, obsolete on last quarter
whorl. Spiral lirae on spire whorls thin, angular, prickly
where cross axials, upper one situated a short distance
below suture, lower one slightly above succeeding
suture. Anterior end of body whorl with 14 spiral lirae,
those on rostrum raised, subequal, flattening out and
becoming more widely set above rostrum, where they
appear as paired furrows.
Pale salmon-coloured, sometimes darker on
periphery of body whorl, ribs and subsutural region
white, salmon-tinged 1in area above suture.
Protoconch papillose, of slightly fewer than 2 whorls,
smooth, apically moderately convex and bearing a
conspicuous brown spot; breadth 0.63-0.70 mm.
Operculum oblong-ovate, yellowish-brown.
Dimensions.
10.6 x 41 mm (holotype), larger paratype (with
juvenile lip) with length 11,2 mm.
Notes.
Immature paratypes of this species show much
similarity to Columbella chuni Thiele, 1925, based on
juvenile material from 404-463 m off Tanzania
(THIELE 1925: 142, pl. 19, fig. 6). Comparison with
syntypes of C. chuni (BMNH 1948.12.10.1-2) shows
that À. blignautae differs in its non-shouldered whorls,
flat-sided, sharper spire and much more elongate base
and aperture. From the three Western Atlantic species
referred here by RADWIN (1978) it differs in its
elongate aperture and more fusiform shape (almost
suggestive of the New World genus Sfrombina Môrch,
1852).
Etymology.
Named after Mrs Tracy Blignaut, assistant to Brian
Hayes.
156
Family OLIVIDAE
Genus Ancillista Iredale, 1936
KILBURN (1993: 372) regarded Ancillaria hasta
Martens, 1902, of the Agulhas Bank, as an atypical
member of this genus. A second South African species
is here added, on the grounds of its large protoconch
and non-ridged columella base. Nevertheless it is
unique within the Ancillinae in its totally smooth,
straight, non-differentiated columellar pillar. When the
body 1s known, this species will probably prove to
belong to an undescribed genus. Terminology after
KILBURN (1977).
Ancillista depontesi n. sp.
Figs 3-5
Type material.
Holotype NMSA V4381/T1532, off Kenton-on-Sea,
101 m, coarse sand and shell debris, dead, Natal
Museum Dredging Programme. Paratypes 1-2, NMSA
V6144/T1553, same data as holotype; paratype 3,
NMSA V6143/T1551, between Great Fish and
Keiskamma River mouths, 100 m, in crayfish trap, with
operculum, B. Hayes. Paratypes 4-6 in B. Hayes
collection, paratype 4, same data as paratype 3;
paratype 5, off Algoa Bay, 100 m, in crayfish trap;
paratype 6, off Port Alfred, 100 m, crayfish trap.
Additional (non-type) material.
"Zululand", 100-200 m, crayfish trap, locality doubtful,
in B. Hayes collection.
Type locality.
Off Kenton-on-Sea (33°55.6'S; 26°440'E), eastern
Algoa Bay, 101 m.
Distribution.
Eastern Agulhas Bank, from off Algoa Bay to the
Keiskamma/Great Fish River area.
Diagnosis.
Shell cuneiform with bluntly rounded apex, rather flat-
sided spire and wide anterior end; base of columella
broad and straight, not twisted, nor defined by an
anterior fasciolar groove; primary spire callus thin,
covering body whorl, slightly indented where covers
suture, and forming a low ridge above suture, without
distinct microscopic granules, although these are
present on the thin secondary callus pad at end of
penultimate whorl and on columella; ancillid band
almost level, defined by shallow grooves: off-white
with a milk-white zone below suture, followed by a
narrow light to dark orange-brown band. Maximum
length 20.4 mm.
KILBURN Four new neogastropods APEX 13(4) 155-160, 20 déc. 1998
Figs 1-2. Anachis (Suturoglypta) blignautae (Columbellidae), n. sp. Holotype NMSA V6140/171548, off Richard's
Bay, Zululand, 600 m, dimensions 10.6 x 4.1 mm. Figs 3-5. Ancillista depontesi (Olividae), n. sp. Figs 3-4. Holotype
NMSA V4381/71532, off Kenton-on-Sea, E. Algoa Bay, 101 m, dimensions 18.2 x 7.6 mm. Fig. 5. Paratype
V6144/T1553, same locality, dimensions 16.3 x 6.3 mm.
157
APEX 13(4): 155-160, 20 déc. 1998
Four new neogastropods
KILBURN
Description.
Shell wedge-shaped with blunt apex, rather straight-
sided spire and broad anterior end, breadth/length 0.39-
0.42; body whorl more convex on left side than on
nght, greatest width of shell at about 0.30 length from
base, aperture/total length 0.44-0.47; apical region
slightiy cyrtoconoid, protoconch obtuse and rather
rounded, spire angle about 30°. Primary spire callus
thin, covering body whorl, slightly impressed at suture,
which it scarcely masks, protoconch exposed; surface
of spire callus glossy, not distinctly microshagreened,
lacking spiral sculpture other than a low angular ridge
of callus immediately above suture; secondary callus
very thin, forming a microshagreened pad at end of
penultimate whorl and covering inner lip.
Aperture cuneiform, gaping basally, widest about
0.25 from anterior end, columella slightly
foreshortened; outer lip thin, in side view evenly
convex, without an ancillid tooth, siphonal notch
deeply, broadly and asymmetrically concave.
Columella not forming a twisted pillar, broad and
straight, with very fine microshagreen sculpture. No
anterior fasciolar groove; inner lip very shallowly and
evenly concave anterorly, straight posteriorly. Ancillid
band very slightly declivous, almost level, demarcated
by a very shallow groove on either side; median zone
with rather coarse growth lines. Termination of
protoconch not demarcated, total number of whorls
about 5.5.
Cream-colour, subsutural margin mulk-white,
followed by a diffuse light to dark orange-brown stripe.
Operculum transparent pale vyellowish, rounded-
trigonal with non-terminal, eccentric nucleus, about
0.45 length of aperture.
Dimensions.
18.2 x 7.6 mm (holotype), 20.4 x 8.4 mm (largest
paratype).
Notes.
Ancillista depontesi bears little resemblance to any of
its congeners, but is superficially most similar to Bullia
ancillaeformis E. A. Smith, 1906, in the Nassariidae.
This resemblance is obviously the result of
convergence, as it differs from the latter in possessing
an ancillid band. From the other Agulhas Bank species,
Ancillista hasta (Martens, 1902), it differs inter alia in
the body whorl being uniformly covered with primary
callus and in the presence of an ancillid band and
groove.
Etymology.
Named in honour of Captain Zeca de Pontes, who first
discovered this unusual species.
Superfamily CONOIDEA
Family DRILLIIDAE
Genus Splendrillia Hedley, 1922
The material of this genus then available from southern
Africa was revised by KILBURN (1988:206-218), who
recorded a total of 8 species. The species described
below is referred to Splendrillia on account of the
restriction of spiral sculpture to the rostrum.
Splendrillia hayesi n. sp.
Figs 6-7
Type material.
Holotype NMSA V6142/T1550; paratype 1, in B.
Hayes collection.
Type locality.
Off Algoa Bay, 100 m, in crayfish pots.
Distribution.
Eastern Agulhas Bank, known only from the type
locality.
Diagnosis.
Claviform (breadth/length 0.36-0.37, aperture/total
length 0.36-0.40), rostrum bent to right, with a strong
fasciole and chink-like umbilicus;, whorls strongly
rounded, upper third concave, flattened below suture;
anal sinus deep, asymmetrically U-shaped, constricted
by thick parietal pad; moderately glossy, axial ribs low,
opisthocline, 9 on body whorl, evanescing below suture
and on base at parietal level, no spiral sculpture except
numerous, weak threads on rostrum; protoconch large
and papillose; uniform white. Maximum length 26.7
mm.
Description.
Shell claviform (breadth/length 0.36-0.37,
aperture/total length 0.36-0.40), of 7 teleoconch
whorls, with blunt apex, body whorl obconical with
moderately short, distinctly oblique, tapering anterior
end, suture moderately shallow, not undulating; whorls
strongly rounded, more angular on early whorls,
periphery just below midwhorl, forming a slight
shoulder on last whorl; upper third of each whorl
concave, flattening out below suture (without a distinct
cord or sulcus); left side of anterior end of body whorl
concave, with a strong fasciole (bending rostrum to
left) and chink-like false umbilicus. Aperture oblong-
pyriform, greatest width at about posterior third,
siphonal canal moderately deep and wide, rather
Figs 6-7. Splendrillia hayesi (Drilliidae), n. sp. Holotype NMSA V6142/T1550, off Algoa Bay, 100 m, dimensions
22.7 x 8.2 mm. Figs 8-9. Turris faleiroi (Turridae), n. sp. Holotype NMSA V6145/11554, off Algoa Bay, 100 m,
dimensions 40.4 x 12.3 mm.
158
KILBURN
straight, termination not dorsally indented. Inner lip
almost straight, with thick callus, edge concave in
parietal region where callus forms a thick posterior
pad, constricting anal sinus. Outer lip chipped in all
types but strongly convex in side view, with deep,
rather asymmetrically U-shaped anal sinus, stromboid
notch evidently very slight.
Surface moderately glossy; sculptured by low axial
ribs only, except for numerous, weak spiral threads on
rostrum;, no definite prelabral varix, growth lines
coarse. Axial ribs opisthocline, in transverse section
angularly rounded, more or less equal to intervals, 9 on
Ist whori, 10-12 on penultimate whorl, becoming
obsolete on last 0,2 of body whorl; ribs obsolete below
suture and at panetal level, appearing as smooth,
oblong nodules on body whori. Uniform white.
Protoconch large and papillose but too worn or
encrusted for details.
Dimensions.
22.7 x 8.2 mm (holotype), 26.7 x 10.3 mm (paratype).
Notes.
Of its known South African congeners, only the much
narrower, salmon-coloured Splendrillia daviesi
Kilburn, 1988, approaches S. hayesi in size. Of Indo-
Pacific taxa the most similar is probably S. solicitata
(Sowerby, 1913) of the Western Pacific, which has
distinctly shouldered axial ribs and faint colour zones.
S. hayesi is superficially similar to another Agulhas
Bank species, Agladrillia ukuminxa Kïiburn, 1988, but
Four new neogastropods
APEX 13(4): 155-160, 20 déc. 1998
that is much smaller, with spiral threads overall and a
straight rostrum.
Family TURRIDAE
Genus Turris Rôding, 1798
The southern African species of this genus were
revised by Kilburn (1983: 552). Much further material
has subsequently been acquired during the Natal
Museum Dredging Programme and will be dealt with
in a future paper. However, the present species 1s
particularly noteworthy in being the first temperate-
water member of the genus known.
Turris faleiroi n. sp.
Figs 8-9
Type material.
Holotype NMSA V6145/T1554: paratype 1, NMSA
V6146/T1555, juvenile; paratypes 2-3 in B. Hayes
collection; all from type locality, in crayfish pots.
Type locality.
Off Algoa Bay, 100 m.
Distribution.
Eastern Agulhas Bank, known only from the type
locality.
159
APEX 13(4) 155-160, 20 déc. 1998
Diagnosis.
Medium-sized, fusiform with aperture/total length
0.34-0.,36, spire orthoconoïd, whorls moderately flat,
shoulder sulcus shallow, subsutural cord broad, with 3-
4 spiral lirae, shoulder cord moderately thin, weakly
crenulate, perniphery of base with 3 strong cords and
weak intermediaries, collabral threads sharp and
crispate, rendering most spiral lirae pliculate; anal sinus
shallow; protoconch large (breadth 1.60-1.75 mm);
reddish-brown, spiral lirae paler, flecked below suture
and elsewhere with pale reddish-brown. Maximum
length 43.5 mm.
Description.
Shell with 11 teleoconch whorls, breadth/length 0.30-
0.31), aperture short (aperture/spire 0.34-0.36), spire
orthoconoïid with flattened whorls and shallow suture;
subsutural cord low, rather 1ll-defined, bearing 3-4
spiral lirae and microscopic spiral threads, crenulated
or pliculated by axial sculpture, shoulder sulcus
shallow. Anal sinus shallow, asymmetrical: shoulder
(sinus) cord as strong as other cords, weakly and
irregularly crenulated. Periphery of base with 3 well-
defined, subequal main cords, each pair separated by 1
intermediary lira flanked by several weaker threads; 2
of these cords are visible on spire whorls (making a
total of 4 cords per whorl), although the lower may be
hidden in suture; 1st teleoconch whorl already with 4
thin lirae, the lower two closer together. Anterior end
of body whorl with 20-25 lirae, those on rostrum close
and even, those above stronger and more wide-set, their
intervals with fine spiral threads. Fine, sharp, crispate
collabral threads overall, crenulating all main lirae.
Light brown with paler main lirae, bearing an
occasional slightly darker reddish-brown fleck,
protoconch light brown. Traces of dull light brown
periostracum retained interstitially.
Protoconch papillose, large and blunt, of about 1.8
whorls, last 0.6 whorls with 8-16 strong, arcuate axial
nbs, with a spiral lira developing a short distance above
suture near termination; breadth 1.60-1.75 mm.
Dimensions.
40.4 x 12.3 mm (holotype), 43.5 x 12.2 mm (largest
paratype, lip damaged).
160
Four new neogastropods
KILBURN
Notes.
Turris faleiroi is very similar to another South African
endemic, Zurris orthopleura Kïilburn, 1983, which
lives somewhat further east on the continental shelf of
Transkei and southern Natal. It differs from 7
orthopleura in its larger protoconch (breadth 1.60-1.75
mm against 1.30-1.50 mm), which is less papillose but
has a more inflated first whorl; the suture in 7° faleiroi
is much deeper, axial threads are much stronger and
sharper, rendering the spirals somewhat crenulate, the
subsutural cord bears 3 distinct spiral lirae, instead of
only fine threads, spiral cords are slightly stronger, the
shoulder sulcus is deeper and the ground colour pale
brown instead of white.
Etymology.
Named after Mr Ginger Faleiro, captain of the crayfish
boat that first discovered this species.
REFERENCES
KILBURN, R. N. 1977. Descriptions of new species of
Amalda and Chilotygma, with a note on the
systematics of Amalda, Ancillus and Ancillista.
Annals of the Natal Museum 23(1): 13-21.
KILBURN, R. N. 1983. Turridae (Mollusca: Gastropoda)
of Southern Africa and Mozambique. Part 1.
Subfamily Turrinae. Annals of the Natal Museum
25(2): 549-585.
KILBURN, R. N. 1988. Turridae of Southern Africa and
Mozambique. Part 4. Subfamilies Drilliinae,
Crassispirinae and Strictispirinae. Annals of the Natal
Museum 29(1): 167-320.
KILBURN, R. N. 1993. Notes on some South African
Ancillinae, with descriptions of five new species of
Amalda. Annals of the Natal Museum 34(2): 369-389.
RADWIN, G. E. 1978. The family Columbellidae in the
western Atlantic. Part IIb. The Pyreninae (continued).
The Veliger 20(4): 328-344.
THIELE, J. 1925. Gastropoda der Deutschen
Tiefsee-Expedition, 1898-1899. II Teil.
Wissenschaftliche Ergebnisse der deutsche
Tiefsee-Expedition auf dem Dampfer Valdivia' 1898-
1899. Jena: G. Fischer. 17(2): 36-382.
TURSCH
Shell model
APEX 13(4): 161-176, 20 déc. 1998
A simple shell model: applications and implications
Bernard TURSCH
Laboratoire de Bio-Écologie, Faculté des Sciences.
Université Libre de Bruxelles, 50 av. F.D. Roosevelt, 1050 Brussels, Belgium.
KEY WORDS, Gastropods, shell, mode!
ABSTRACT, A simple computer-assisted, non-mathematical procedure for emulating sagittal
sections of Gastropod shells is described. Examples illustrate that the final “shell” shapes can largely
be predicted from the values of the construction parameters. The problem of the meaning of
traditional descriptions of the shape of shells is briefly addressed.
1. INTRODUCTION
Coiled shells are the hallmark of most living Molluscs
and all Brachiopods, amounting to about half of all
non-arthropod invertebrates. They also constitute a
large proportion of all fossils. The fascination long
exerted on biologists, mathematicians and other artists
by the regular shapes of shells is reflected in a copious
literature and an abundant iconography. As it could be
expected, many mathematical models have been
proposed to explain or imitate the growth of coiled
shells. Most of these models have been reviewed by
MEINHARDT (1995) and STONE (1996). For many years,
the standard tool for the geometrical analysis of coiled
shells has been the model developed by the eminent
palaeontologist D.M. Raup in a series of papers
culminating in his well-known 1966 synthesis.
À simple, operational modei of coiled shells has
been recently developed (TURSCH, 1997a). The model
was intended as a probe for biological studies rather
than for realistic simulation of specialised structures, so
it could be kept very easy. It has several advantages
over other shell models. Amongst others, it rests upon
independent parameters and can simulate shells with
non-isometric growth (for instance Gastropods with
concave or convex spires) without having to postulate
ad hoc changes in the shell parameters (which amounts
to make constants vary). The basic shape of the shell
(this does not account for spines, sculpture, etc.) is
entirely determined by the construction parameters.
One short paragraph in the original paper stated that
the outcomes of the construction are largely predictable
by comparing the values of the parameters. Detailed
examples will now be given.
Some conchologists have been made insensitive to
the joys of mathematics. Yet they can easily produce
rough simulations of sagittal shell sections without
using any equation at all, by using a small computer
equipped with one of the many drawing programs now
in common use. The step by step procedure (very
summarily outlined in TURSCH, 1997a) will be
described here. It is particularly suited for simulating
the shells of multi-coiled Gastropods.
À computer program that automatically generates
“shells” can easily be derived from the model. If one
aims at the mass production of shell models, the use of
such à program will save several minutes on every
construction. If one aims at understanding the role of
the individual parameters and appreciating how these
parameters do interact, then the step by step, hands-on
procedure is certainly more informative.
Understanding shell parameters can be of
importance for evaluating the descriptions of the shape
of shells, which are at the very foundations of mollusc
taxonomy. Let us consider the two shells depicted in
Fig. 1. The obvious difference in their aspect would
ordinarily be described by listing differences in the
states of a series of traditional shell characters. These
may be the general outline of the shell, the height of the
spire, the shape and orientation of the aperture, the
convexity of the whorls, etc. In works of taxonomy, the
question of whether the characters in this list are
independent of each other or are not is very rarely
raised, if ever. It might be instructive to see how these
different traditional characters relate to differences in
shell parameters.
This paper is about shapes and relies heavily on
illustrations. For the study of shapes, one drawing
speaks better than a thousand words, a bunch of
equations or a few pages of computer program listing.
Figure 1. The problem of shell description. How do
these two shells differ? (see text $ 1).
161
APEX 13(4): 161-176, 20 déc. 1998
}
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Shell model
TURSCH
Figure 2. The shell model: general principle (see text $ 2.1).
2. THE MODEL
2.1. Generalities
As in most other shell models, the "shell" 1s the surface
of revolution produced by a regularly growing
generating curve (the shell aperture) effecting a helico-
spiral motion along an axis (the coiling axis). The
generating curve K, is, as usual, taken to be an ellipse
because the aperture of most shells can be
approximated by (or inscribed in) this shape.
To simulate the sagittal section of a shell (such as
the shell in Fig. 2, a) one first has to position in relation
to a coiling axis a starting ellipse K, of suitable shape
and size (Fig. 2, b).
One then determines where the centre of the
generating curve K, will be located at each subsequent
half-volution (Fig. 2, c). C, is the centre at the start and
Cos, C1, C15, C2, .…., C, are the centres after 0,5, 1, 15;
2, …, n revolutions. The position of these successive
centres are found by building the successive rectangles
"0.5", "1", "1.5", etc. They are all simply derived (by
the use of appropriate parameters) from an essential
element in the starting configuration: rectangle "O0"
(darkened in Fig. 2, c and d). Obtaining rectangle "0"
will be explained in detail in $ 3.1.1.
The starting figure K, is then "grown" by an
appropriate factor to obtain Kss, K1, Kiss, K2, etc., each
of which is the placed on its calculated centre Cos, C1,
C 5: CG (Fig. 2 d).
If so desired, sutures can be drawn and the aspect of
the whorls can be simulated by joining the edges of
Kos, K1, Kis, K>, etc. with appropriate lines (Fig. 2, €).
The suitably positioned ellipse K;, the three points
Co, C1, C> and three growth parameters do completely
determine all the construction, no matter the number of
whorls.
162
2.2. Parameters
The parameters of the model have been defined in
TURSCH (1997a). This has to be repeated here, to make
the construction procedure comprehensible.
coiling
axis
4 ;
| 10 es;
Figure 3. Parameters for shaping and positioning the
generating curve Ko (see text $ 2.2.2).
2.2.1. Parameters determining the starting
conditions
The size and proportions of the starting ellipse K, (see
Fig. 3) are determined by its smallest diameter w, (here
always equal to 1) and its ellipticity e (the ratio of its
longer axis to the shorter).
The spatial orientation of the ellipse K, in relation
with the coiling axis is described in the complete model
(see TURSCH, 1997a) by three angular parameters ©, f
and à. In the simplified, rough simulation presented
here, the generating ellipse K, is always co-planar with
the axis, so parameters f and à will bc neglected.
Parameter « is the angle of the long axis oi Ko with the
Coiling axis.
TURSCH
Parameter q is defined as:
q=ro/(ho/2) so ro= q.(ho/2)
Particular case: if angle &« = 0 then
ho=wo and ro = q.(Wwy2)
If angle à = 0 and if K, is tangent to the coiling axis (a
common case) then ro = (w./2) and g= 1.
2.2.2. Parameters positioning the centre after one
volution
Positioning the centre C; amounts to determining &
and r; (see Fig. 4). Parameter p has been defined as
Pp=do/(voy/2) so: d5= p.(vo2)
Particular case: if angle « = O then
Vo — €.Wo and do D: (ewo/2)
Parameter À, (the rate of Radial expansion) has been
defined as
R,=n/ro so n=A"
Parameter À applies to all subsequent whorls, so
Re To Feet (1)
2.2.3. Parameters positioning the centre after two
volutions
Positioning point C:
determining d, and r:.
Parameter £ (the rate of Longitudinal expansion) has
been defined as
L= d\ /d so d = £.d
This parameter applies to all subsequent whorls, so
(see Fig. 4) amounts to
L=d/d4=d,/d,) (2)
r depends on parameter À defined here above:
r=kR"n
2.2.4. Growth of the generating curve
This amounts to determining w,, the diameter after one
revolution.
The growth of the generating curve after one volution
determines parameter W (the rate of Whorl expansion)
W=wW/w
One will notice that W is the same as Raup's parameter
W. This parameter applies to all subsequent whorls, so
W=wWm/w=w,/Ww,: (3)
2.2.5. Subsequent volutions
Each subsequent centre C, is placed in relation to the
preceding centre C,;1 by direct application of
parameters À, and £ [see expressions (1) and (2)]. The
size of each subsequent motive K, 1s simply that of the
previous motive K,, multiplied by parameter W [see
expression (3)].
2.2.6. Remarks
The internal definition of £ (the only originality in this
otherwise obvious model) allows one to dodge the
problem of having to select a point of origin for the
helico-spiral. The position of this point in relation to Ko
determines much of the shape of the resulting surface
of revolution, a difficulty that has plagued previous
models.
Shell model
APEX 13(4): 161-176, 20 déc. 1998
Figure 4. Construction: parameters for developing shell
whorls (see text 8 2.2.1).
The parameters of the model are of two very
distinct kinds (see TURSCH 1997a). Parameters q, p, e,
and « are fixed initial conditions, and it is tempting to
speculate that they reflect an embryonic répertoire (see
TURSCH 1997a). Parameter p only sets the pitch of the
first volution. Parameter q is defined from an initial
distance r, and is useful for model construction and
analysis convenience. In contrast, parameters W. R,
and £ are expansion rates. They just selectively
amplify the starting parameters during, growth, as long
as ñ (the number of volutions) has not reached its final
value.
3. APPLICATIONS
Drawing program requirements. The program should
be able to draw lines, rectangles, ellipses and circles. It
should also be able to group, move, rotate, mirror and
scale objects (by stretching vertically and horizontally)
by a given percentage. Most of the recent drawing
programs allow these operations.
Graphic conventions. The step by step graphic
constructions are made mostly by stretching and
moving selected elements. In each step, the copy of a
starting element (thick lines, light shading) is stretched
horizontally by x% and vertically by y% (indicated by
H= x%;, V= y%). It is then moved as indicated by
arrows to yield a resulting element (very thick lines,
dark shading). This is often the starting element for the
next step. For typographic facility, square roots are
indicated in the text as: p°”.
163
APEX 13(4): 161-176, 20 déc. 1998 Shell model TURSCH
2e
PO
Fe ne" RE.
a D
H=
(140 %)
L
(150 %)
(130 %) (130 %)
Parameters
6=200 : @=20 :"q= 15 101 p=1250 W= 1302 = 1/40" LM SD MNES
Figure 6. Construction of sagittal half sections (see text $& 3.2).
164
TURSCH
3.1. Construction: starting elements
AII constructions require the same first steps: the
positioning of the starting ellipse K; and the
construction of rectangle "0".
3.1.1. Starting ellipse
Parameters needed: e, q and «. Let us take as example:
EC 2Na- So 20,
a. Draw two crossed lines (see Fig. 5, a). From their
intersection as a centre, draw a circle of diameter w
(this has necessarily an ellipticity e = 1). With the
command "group" (or similar) associate the lines with
the circle into one single picture (the position of the
centre will be needed to allow accurate positioning
during the remainder of the construction). w, will be
the length unit.
b. This figure is now stretched vertically by 200 % (in
order to obtain an ellipse with the desired ellipticity e =
2) (see Fig. 5, b).
c. Rotate the ellipse by an angle & (in this case 20°)
(see Fig. 5, c).
d. Draw the coiling axis and any line perpendicular to
the axis (see Fig. 5, d). Build segment a.
e. Stretch horizontally a copy of segment a by 150 %
(because qg = 1.5) to obtain segment b, which is then
placed as shown (see Fig. 5, e).
f. Position the ellipse at a distance b from the coiling
axis (see Fig. 5, f). The generating curve K, is now
fully positioned, with its centre Co, marked by
intersecting lines.
g. Erase all unnecessary features. Draw rectangle "O0"
(see Fig. 5, g). This will be the stepping stone for the
remainder of the construction.
3.1.2. Particular cases
a. The construction is simplified if « = 0 (ellipse
parallel to the axis). Segment a (see Fig. 5, h) is now
the half diameter w,/2 of the ellipse. The next two steps
(see Fig. 5, 1 and j) are straightforward.
b. Things are especially simple if à = 0 and g = 1 (a
very common case). AIl one has to do is then to bring
directly the ellipse tangent to the axis (see Fig. 5, k)
and draw rectangle "0" (see Fig. 5, 1).
c. One will note that if e = 1 then « is indeterminate
(rotating a circle by any amount vields the same circle).
TIPS: Make the starting ellipse small enough because it
might grow very much (your computer program can
"zoom" on small features). Most shells can be
simulated by placing the starting ellipse tangent to (or
very close to) the axis.
3.2. Construction: sagittal half sections
These constructions are very fast and extremely simple
because no calculation at all is needed. Sagittal half-
sections most often contain enough information to
grasp the final shape of the whole “shell”. All
parameters are set in advance. The recipe is 1llustrated
step by step in Fig. 6.
Shell model
APEX 13(4): 161-176, 20 déc. 1998
3.2.1. Starting elements
The starting elements (Fig. 6, a) are obtained as
described in $ 3.1, illustrated in Fi S. In this example:
eq Sa 20;
3.2.2. Centre after first volution.
Parameters needed: ‘R and p. In this example À = 1.4,
DADS;
The position of centre C; (Fig. 6, b) is found by
stretching a copy of rectangle "/" horizontally by 140%
(because À = 1.40) and vertically by 125% (because p
= 1.25).
3.2.3. Centre after second volution.
Parameters needed: ‘R and £. In this example: À =
JAOPLEUrS;:
The position of centre C; (Fig. 6, c) is found by
stretching a copy of rectangle "/" horizontally by 140%
(because À = 1.40) and vertically by 150% (because L
= 1.5). This new rectangle (rectangle "2") 1s placed as
shown in Fig. 6, c. The remainder of the positioning of
the subsequent centres is now repetitive.
3.2.4. Centres of subsequent volutions
Parameters needed: À and £, as above.
The position of centre C; (Fig. 6, d) is found by
stretching a copy of rectangle "2" horizontally by 140%
(because À = 1.40) and vertically by 150% (because L
= 1.5). This new rectangle (rectangle "3") is then
placed as shown. For a "shell" with n volutions, the
same procedure 1s repeated until one obtains rectangle
"n", determining the position of centre Cn.
3.2.5. Generating curve after one volution
Parameter needed: 'W. In this example: W= 1.30.
Figure K,; is obtained (see Fig. 6, c) by stretching a
copy of the starting ellipse K; horizontally and
vertically by 130% (because #/ = 1.30). This new
figure is then placed with its centre (marked with
intersecting lines) exactly at point C;. The "growth"
and the positioning of the generating curve at the
subsequent volutions are now repetitive
3.2.6. Generating curve at subsequent volutions
Parameter needed: W, as above.
Figure K; is obtained (see Fig. 6, f) by stretching a
copy of the starting ellipse K; horizontally and
vertically by 130% (because #/ = 1.30). This new
figure 1s then placed with its centre (marked with
intersecting lines) exactly at point C:. The "growth"
and the positioning of the generating curve at the
subsequent volutions is now repetitive (see Fig. 6, g).
For a "shell" with » volutions, the same procedure 1s
repeated until one obtains ellipse K,, centred on C,.
3.3. Construction: sagittal full sect:ons
AIl parameters are set in advance. The procedure now
entails the construction of the “shell” at each half-
volution. Two steps do require simple transformations
of the parameters.
165
APEX 13(4): 161-176, 20 déc. 1998
Shell model
TURSCH
(140 %) (84.53 %)
V= p V=1/(1+Vp) H=VR
(125 %) (47.21 %) (118.3 %)
V=£I/(1+\V£)
ï (67 41 %)
(1225 %)
= VW = VW = V
(114 %) (114 %) (114 %)
V= VW = VW V= VW
(114 %) (114 %)
Parameters: 6 =200 ; @=20 ; q=15 ; pP=1.25 ;
Derived values: 1/(1+\/p)=0.4721 ; VW=1140 ;
W=130::R=AMIEML=150%N-3
VR =1183 ; 1/VR =-0.8453 ; VL =0.6441 ; L/(1+ VL) = 0.6441
Figure 7. Construction of sagittal full sections (see text & 3.3).
166
TURSCH
3.3.1. Starting elements
As for sagittal half-sections ($ 3.2), the starting
elements (Fig. 7, a) are obtained as described in $ 3.1
and illustrated in Fig. 5. In this example: e = 2: q = 1.5:
a = 20.
3.3.2. Centre after first volution
This step is the same as for sagittal half-sections (see $
827)
Parameters needed: R and p. In this example À = 1.4,
p— 125
The position of centre C; (Fig. 7, b) is found by
stretching a copy of rectangle "/" horizontally by 140%
(because À = 1.40) and vertically by 125% (because p
= 1.25).
3.3.3. Centre after 0.5 volution
Parameters needed: ‘R and p. One has to calculate the
values of °° (here: 1.183) and 1/(1+ p°*) (here:
0.4721).
The position of centre Co: is found by stretching a
copy of rectangle "0" horizontally by 118.3% (because
R°° = 1.183) and vertically by 47.21% [because 1/(1+
p°°) = 0.4721]. This new rectangle (rectangle "O.5") is
placed as shown in Fig. 7, c.
3.3.4. Centre after 1.5 volution
Parameters needed: R and £. One has to use R°° (in
this example: 1.183) and £/(1+£L°*) (here: 0.6741).
The position of centre C; is found by stretching a
copy of rectangle "7" horizontally by 118.3% (because
R°° = 1.183) and vertically by 47.21% [because L/(1+
£°$) =0.4721]. This new rectangle (rectangle "/.5") is
placed as shown in Fig. 7, d.
3.3.5. Centre after two volutions
Parameters needed: R and £. One has to use °° (in
this example: 1.183) and £°*) (here: 1.225).
The position of centre C; is found by stretching a copy
of rectangle "/.5" horizontally by 118.3% (because °°
= 1.183) and vertically by 122.5% (because L°°=
1.225). This new rectangle (rectangle "2") is placed as
shown in Fig. 7,e.
3.3.6. Centres of subsequent volutions
Parameters needed: R°* and L°*, as above.
The position of centre C;5 (Fig. 7, f) is found by
stretching a copy of rectangle "2" horizontally by
118.3% (because R°° = 1.183) and vertically by
122.5% (because L°*= 1.225). This new rectangle
(rectangle "2.5") 1s placed as shown. For a "shell" with
n volutions, the same procedure is repeated until one
obtains rectangle "n", determining the position of
centre C, (see Fig. 7, g).
3.3.7. Generating curve after 0.5 volution
Parameter needed: W°*. In this example: W = 1.30
and W°*= 1.14.
Figure K, : is obtained (see Fig. 7, h) by rotating a copy
of the starting ellipse K, by an angle -«, then stretching
it horizontally and vertically by 114% (because #4 * =
1.14). This new figure is then placed with its centre
Shell model
APEX 13(4): 161-176, 20 déc. 1998
(marked with intersecting lines) exactly at point Cos.
The "growth" and the positioning of the generating
curve at the subsequent volutions are now repetitive.
3.3.8. Generating curve at subsequent volutions
Parameter needed: W°*, as above.
Figure K, is obtained (see Fig. 7, 1) by “rotating a copy
of the ellipse Koss by an angle -«, then stretching it
horizontally and vertically by 114% (because W ° =
1.14). This new figure is then placed with its centre
(marked with intersecting lines) exactly at point C1.
The "growth" and the positioning of the generating
curve at the subsequent volutions is now repetitive (see
Fig. 7, j). For a "shell" with » volutions, the same
procedure is repeated untii one obtains ellipse K,,
centredonC,.
The procedure might look more difficult than it really
is. With a little practice, once the derived values have
been established, steps a to k (in Fig. 7) are easily
effected in less than 5 minutes.
3.3.9. Final image
The final image (Fig. 7, k) can now be made up by
masking hidden parts, drawing sutures and delineating
the shape of whorls (for instance as in Fig. 7, 1, m or n).
Whorli resorption occurs in many Gastropods.
According to the desired type of model, one can elect
to have the “aperture” mask the previous whorl or not.
e=4 e=150
= 110 a=0
D = 406 D =060
= 4 =
‘W= 5.00 W= 150
R= 5.00 R=150
£=5.90 L=180
n=2 n=5
a
e=8 e = 150
a= 15 a=0
g=i g=1
p=095 p = 2.00
W= 150 ‘W= 120
R=150 R= 120
L=1.45 L£=120
n=7 n=9
3H 200 R ©
TL CL 1
Figure 8. Construction: examples of applications (see
text $ 3.3.9).
167
APEX 13(4): 161-176, 20 déc. 1998
e=2
e=var £=1.50
g=100 ‘W-=-130 : x
p=125 ÆR=140 e varies
Shell model
A Æ
g=1.50
e=2 £L=1.50
g= var W= 1.30
TURSCH
D varies, | P-125 ÆR-140
L=180
g=15 W=130
| L varies
Figure 9. Shape variations due to changes in a single parameter (see text $ 4.1).
Rather realistic renditions of many existing shells are
easily produced by the graphic construction described
here above (see Fig. 8, illustrating a few familiar
cases). For even more realism, the shape of the starting
ellipse could be modified, for instance by adding or
substracting suitable features. With so many variables,
it would take a very long time to produce a given
“shell” by experimenting with arbitrary combinations
of parameters. The task is very much simplified 1f one
understands how each individual parameter acts and
how given combinations of parameters do affect the
final shape.
168
4. THE CONTROL OF SHAPE
Easy “rules of construction” can be deduced from the
model. Some are given here under as examples. Many
more could be found by an interested reader.
4.1. Effect of individual parameters
Examples of the changes resulting from the variation of
individual parameters are shown in Fig. 9. The effect of
the expansion parameters W. À, and £ are quite
predictable. One will notice that the final shape is
extremely dependent from the initial conditions g, p
and e.
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world in search for the most beautiful specimens. Not only with the
aim of perfecting your collections, but also to gain more knowledge
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85, rue des Coteaux Fleuris - 83200 TOULON
Tél. 04 94 92 96 21 - Fax 04 94 22 97 46 Gauguini
TURSCH
e = var £=120
g=100 ‘W=1.50
p=150 R=150
Shell model
APEx 13(4): 161-176, 20 déc. 1998
Figure 11. Non-isometric growth without change in shell parameters. The shape of the shell Varies with the number
of whoris n. (see text S 4.1).
35b 200 ®
Il Ma Woo
<D-N
7
CES
nou
ON = —
8
Figure 12. Non-isometric growth by abrupt change of a shell parameter. The construction of shell a yields shell b if
at whorl 2.5 (marked by an arrow) one changes the value of W from 1.5 to 1.2 (see text $ 41).
Fig. 10 shows that e (the ellipticity of the
generating curve) influences not only the shape of the
body whorl but also the shape of the spire. Some more
dramatic effects of parameter e will be shown in $ 43.
The shape of many shells (most shells, according to
VERMEL 1993) does vary during growth. In contrast to
others, this model can produce “shells” with non-
isometric growth (see Fig. 11) without having to
modify progressively the values of parameters. It is
thus quite important to specify a value for » (the
number of whorls).
Abrupt changes in the value of parameters do
happen during the growth of some real shells (for an
example, see TURSCH 1997b). This can of course be
easily emulated in this step by step procedure by
modifying any of the expansion parameters W, R, or L
at any desired point of the construction (for an
example, see Fig. 12).
169
APEX 13(4): 161-176, 20 déc. 1998 Shell model TURSCH
all models: e =2 ; &=20 : q=13 ; R=150 ; n=4
= 070 P = 090
‘W = 1 40 ‘W= 1.40
£L=170 Gen70
/
te
Figure 13. General features of the shell are affected by individual variations of several parameters. For instance, all
other parameters kept constant, the length of a given shell (a) is modified by a change of L (b), of p (c) or W(dj.
(see text $ 4.2).
R<L R=L
Figure 14. The relation of R to L determines the alignment of the centres (see text $ 4.2.1).
all models: Æ=£ = 150
W=135
R=L>W R=L=W R=L<W
spire convex spire conical spire concave
Figure 15. If R = Lthen the value of W determines the shape ofthe spire (see text & 4.2.2).
General features of the shell are affected by of p, of ‘W or L will affect the total length of the
individual variations of several parameters. For “shell” (see Fig. 13). In the same way, the diameter (at
instance, the total length depends on both the rate at any moment of growth) depends on both the growth
which the centre of the aperture moves "down" the rate of the aperture (W) and the rate at which the
coiling axis (the compounded effects of p and £) and centre of the aperture moves away from the coiling axis
the growth rate of the aperture (#/). Therefore, all (Æ).
other parameters being the same, individual variations
170
TURSCH
CF \
BEA DA
e=2 e=2
JE g=i
p=070 p = 1.50
‘W= 150 ‘W= 150
R=150 R= 150
L=150 L=N20
n=4 n=4
Shell model
APEX 13(4): 161-176, 20 déc. 1998
O NN
—
[®)]
3 b}3 200 ©
il
82
Bb =
œ
[e)
Figure 16. |f q = 1 and W= Rthen all the whoris are tangent to the coiling axis (see text $ 4.2.2).
e=1 e
œ=0 œ
qg=13 q
P = 2.49 p
‘W=300 y
R=150 R
£L=170 JE
n=2 n
NE
Wu un Te Wu Ou HN
Figure 17.IfW> R then fabricational problems may occur (see text $ 4.2.2).
4.2. Effect of combinations of parameters
By their definition, all the parameters are completely
independent from each other (this was not the case of
the parameters in the classical model of RAUP 1966).
However, the shape of the final "shell" depends very
much on the interaction of these independent
parameters.
4.2.1. Parameters À, £ and W
If À = L then, in sagittal view, all the centres are
aligned on a straight line. The revolution of the centre
of the generating curve takes place on a conical surface
(see Fig. 14, b). If À < £ the surface of revolution of
the centres will be convex (see Fig. 14, a), If R > Lit
will be concave. (see Fig. 14, c). Note: these relations
determine only the positions of the centres, not the
outline of the shell.
If R = L = ‘W then growth will be isometric.
leading to shells with true conical spires (see Fig. 15,
b). If R = W + L then growth will be non-isometric,
the shape of the shell varying during growth (see $
4.1). IR =Z£L > W the spire will be convex (Fig. 15.
a). IFR = L <‘Wthe spire will be concave (Fig. 15, c).
4.2.2. Parameter q
If g = 1 then the generating curve K, 1s tangent to the
coiling axis (see the definition of q).
If q = 1 and W= & then ail the whorls are tangent
to the coiling axis, whatever the values of the other
parameters. This is a very common case in real
Gastropods, as illustrated by the few examples in Fig.
16.
If q = 1 and W = R = L then all the whorls are
tangent to the coiling axis and the shell is conispiral
(Fig. 16, a).
If W > R and if the number of volutions is not
limited, the whorls will ineluctably increase beyond the
coiling axis. If g = 1, this overlap will occur right at the
start of the construction. If g > 1 it may happen either
at the start (Fig. 17, a) or after a few volutions (Fig. 17,
b), depending on the values of the other parameters.
This does not necessarily constitute an insurmountable
fabricational constraint. In real shells, the problem may
be solved in various ways, for instance by the
resorption of previous whorls or by changes in the
angular parameter f (not treated here, but see TÜURSCH
1997a).
171
TURSCH Shell model APEX 13(4): 161-176, 20 déc. 1998
5
all models: e=2 ; g=150 ; W=120 ; R=150 ; n
a
PE PE) \
at) dl)
ë Lre SA
œ= 20 œ=0
p=0 p=0
œ= 20 a&=0
p=0 p=0
L=0 L=0
a
Figure 19. Dextral end sinistral shells (see text $ 4.2.4).
original
CA CA A
CA j
A € A is
EDS b
a |
as, CT <
D NS RE
V=25% V=50% V=75% V=100% V=125% V=200% V=100%
size 100 % size 100 % size 100 % size 100 % size 80 % size 50% size 100 %
Original image: € = 1.50 ,@=0 ;, q=1 ;, P=1.50 ; W = 1.20 RU 20 EN 20
Figure 20. The stretching of whole images. The case a = 0 (see text & 4.3).
original
V=150 %
size 667 %
V=125%
size 80 %
V=100%
size 100 %
V = 50 % V=75%
size 200 % size 133,3 %
Original image: € = 8.00, @=15; q=1 ; P=0S,5 ,; W= 1.50 : R=1.50 ; L=145; Nn=7
Figure 21. The stretching of whole images. The case a z 0 (see text $ 4.3).
172
TURSCH
vero
il Wu ou
,0+0N
3bt}>
Shell model
APEX 13(4): 161-176, 20 déc. 1998
LB
—
a
O © O =
œ
I
on
3x 20€ R®
nn
PIN
SIC
ON
Wu un
Figure 22. The problems of the suture (see text $ 5.1).
4.2.3. Parameter p
If p = 0 then L is indeterminate (its value is irrelevant
for the construction). All the centres are located in the
same plane, perpendicular to the coiling axis (Fig. 18,
a).
If p = 0 and à = O0 then the “shell” is isostrophic
(has a plane of symmetry) (Fig. 18, b). Note that the
word “planispiral” has been avoided here, as it can be
taken in different meanings (see COX 1955, ARNOLD
1965).
p ='W -1 and W= £L and « = 0 is the condition for
the “shell” to be discoidal (Fig. 18, c).
4.2.4. Dextral and sinistral “shells”
The observant reader will have noticed that the model
does not specify the direction of coiling. Both dextral
and sinistral “shells” can be obtained from the same
construction (see Fig. 19). Sinistral shells (of entirely
different nature) can be obtained by assigning negative
values to parameter p or to parameter £. Note: one has
then to take the negative square root of the absolute
value.
4.3. Modification of completed models
Once a model has been completed, it is easy to modify
its shape by stretching the whole image (all parts
having been linked into one single image by using the
command ‘“group'). This generates very rapidly
“shells” of various shapes. But what is one then really
doing?
If &« = O in the original image, then vertical or
horizontal stretching modifies only parameter e. An
example of related images obtained by vertical
stretching is given in Fig. 20. The magnitude of the
observed changes in shape fully confirms the
conclusions of $ 4.1. Stretching “shells” does of course
change their sizes. Many will have to be reduced or
enlarged accordingly, to allow better comparison of
shapes.
If &« # O in the original image, then vertical or
horizontal stretching does modify the value of both
parameters e and &, as shown in Fig. 21.
5. IMPLICATIONS
5.1. Suture
The suture has been often used in shell morphometry
because it is mostly easy to observe and lends itself
well to a variety of measurements. However, the suture
is a feature of much more complex nature than
conchologists generally assume.
The suture is the locus of the outermost points
belonging to two consecutive whorls. Determining the
equation of the suture in terms of shell parameters 1s far
from being elementary. Conversely, attempting to
deduce the shell parameters from the suture would be
extremely difficult (1f possible at all).
Careful examination of sutures can nevertheless
give most useful information. Abrupt changes in the
aspect of the suture often indicate abrupt changes in
parameters (for an example, see TURSCH 1997b: 98).
The example depicted in Fig. 22, a shows that the
suture does not necessarily describe a regular helico-
spiral: it starts by going "down" then goes "up" (this
condition, although uncommon, is met in some real
shells with a sunken spire, such as Oliva concavospira
Sowerby, 1914). The revolution surface on which the
suture is inscribed is also not easily deduced from the
surface of revolution of the centres or even from the
profile of the spire. This can be seen on the example of
Fig. 22, b. In this sagittal section, the suture goes
"down" the axis while the spire goes "up".
Small differences in shell parameters can produce
large differences in the aspect of the suture and more
work is definitely needed to clarify the properties of
this familiar shell feature.
173
APEX 13(4): 161-176, 20 déc. 1998
Shell model
TURSCH
CRC
ON O =
un
3LS 200 R ©
a
Wu un
D Oo
Figure 23. Two examples of “impossible shells” (see text $ 5.2).
N
ou
SLR 2020 R ®
Wu nu Mal WOW
M , 200
Won Mall HW HN
ND
[2]
[e]
3 HS 202 R ©
Figure 24. Shell parameters vs. traditional characters of shape. Variation of one single parameter (see text $ 5.3).
all models: e=2 ;,&G=0 ; n=4
qg=2122 q= 130
P = 1063 p = 1.30
W= 1563 ‘W= 1.60
R= 1465 R= 150
L= 1402 L=130
q=130 q = 130
p = 130 P = 1063
W= 1.60 W= 160
R=150 R=150
L= 1.402 BAS
Figure 25. Shell parameters vs. traditional characters of shape. Variation of several parameters (see text 8 5.3).
5.2. “Impossible shells”
Besides imitating known shells, the model can also
produce "“shells” that we can not (not yet? not
anymore?) have in our collections. Many strange
shapes are possible and only two examples will be
given here. Some of these constructions meet obvious
fabricational problems (for instance the “shell” in Fig.
23, b), some others seem perfectly feasible (see Fig. 23,
a).
174
Accumulating a collection of such “impossible
shells” is amusing but is not only a game. It constitutes
an excellent tool for finding and maybe explaining the
"forbidden avenues" of evolution in the ‘“shell
morphospace” (this 1s the set of all possible outcomes
from a given geometrical/mathematical model). The
interest of this classic problem in evolutionary biology
has been recently emphasised by DAWKINS (1996).
TURSCH
5.3. Shell parameters vs. traditional characters
of shape
The basic shape of shells (and of their parts) is usually
described by a series of traditional characters (general
outline of the shell, height of the spire, shape and
orientation of the aperture, convexity of the whorls,
etc.). The correlation between shell parameters and the
conventional shell descriptions raises a number of
questions.
Example A. “Shells” à and f in Fig. 20 are exactly the
same as the shells depicted in Fig. 1. On the one hand,
these shells have a completely different aspect,
reflected by ïarge differences in many traditional
characters of shape. On the other hand, the two shells
are very closely related in terms of shell parameters.
They differ only by parameter e, as can be seen in Fig.
24 where all their parameters of the two shells are now
given.
Example B. Conventional descriptions of the two
closely matching “shells” à and b in Fig. 25 would be
extremely similar, yet these two “shells” differ by the
values of no less than five parameters. The smallness of
the variation of each parameter does not justify the
observed similarity. Let us modify shell b by changing
only parameter Z by the same amount. One then
obtains shell c, of noticeably different shape (see Fig.
25). Modifying only parameter q leads to shell d, of
quite different aspect. The similarity is due to another
cause: the effects of the variations in individual
parameters nearly cancel each other. In real shells, this
would be a nice case of convergence (possibly a case of
sibhing species).
Example A raises an immediate question. Do the
different traditional shell characters really represent
distinct characters? Example B shows that the
traditional descriptors of basic shape do not necessarily
reflect differences in shell parameters.
Example A shows that the conventional characters
of shape are certainly correlated. All are entirely
determined by the parameters of the model. All can
change simultaneously by modifying one single
parameter. Traditional descriptors of basic shape only
appear to be independent. This illusion is simply due to
the reductionistic way by which we describe a complex
structure. We proceed by dividing it in arbitrary,
smaller parts then describing these parts in succession.
The shell parameters being completely independent.
one could be tempted to consider that each of them is a
shell character. This would raise a serious problem.
Indeed, we would then be compelled to consider that
the very different shells a and b are more closely
related than the very similar shells b and c. Fortunately,
this does not happen. Shell parameters do not satisfy
the conditions required for characters measuring
phyletic similarity. They cannot be absent (thus
precluding evolutionary novelty); there are no
“primitive” and no “derived” parameters.
Shell model
APEX 13(4): 161-176, 20 déc. 1998
The very fact that we can (most often) recognise
species by their shells establishes that the shell
parameters, albeit mathematically independent, are
biologically correlated. So there is no “description vs.
parameters” paradox if one considers that it is the
whole set of shell parameters that constitutes one
single, numerical shell character. This holistic approach
of shells reminds of the notion of “morphological
integration” of NEMESCHKAL (1991).
It is the very same shell character that conventional
descriptions attempt to convey (this time with words
instead of figures). If the growth of the shell 1s regular
(with constant parameters) then the whole set of the
many traditional shell “characters” describing the basic
shape of the shell and of its parts constitutes one single
character.
It is not suggested that the whole set of shell
parameters 1s controlled by one single gene! Most
probably, these parameters do not even exist in nature
as separate entities. They are parts of a model that
describes the growth, not of natural law that causes à
particular type of growth.
5.4. Deriving parameters from real shells
This paper concerns the building of conceptual ”shells”
from a set of predetermined parameters. What does it
imply about the reverse operation: deriving parameters
from real shells?
In the simple case of regular growth, the minimum
requirements for finding all the parameters are: the
correct positioning of the coiling axis, the
determination of the co-ordinates of the centres at least
at 3 accurately determined positions, the determination
of the increase of the generating surface between at
least at 3 accurately determined positions.
These very simple requirements are fraught with
problems because small experimental errors in
measurements may lead to serious discrepancies. A
reliable, accurate method for exact positioning of the
axis has yet to be published. Determination of the
position of the centres is anything but evident,
especially if the generating curve is not a true ellipse (it
rarely is). Further problems arise because, in contrast to
real shells, the theoretical shell model is an immaterial
surface, without any thickness. One should also note
that the same difficulties will be met with all other
helico-spiral shell models.
Similar shells may differ by a number of parameters
(see $ 5.3.B), so really accurate determination of their
values seems a priori quite difficult. To estimate shell
parameters, graphic simulations are possibly more
operational than shell measurements.
Acknowledgements. I gratefully acknowledge the
support given by the FNRS and by BIOTEC, S.A. to this
laboratory. I thank Christian Van Osselaer for helpful
criticism.
APEX 13(4): 161-176, 20 déc. 1998 Shell model TURSCH
6. REFERENCES
ARNOLD, W.H. 1965. A glossary of a thousand-and-
one terms used in conchology. Feliger 7
(Supplement): 1-50.
Cox, L.R. 1955. Observations on Gastropod
descriptive terminology. Proc. Malac. Soc. 31: 190-
202.
DAWKINS, R. 1996. Climbing mount improbable.
Viking, England.
MEINHARDT, H. 1995. The algorithmical beauty of sea
shells. Spnnger, Berlin.
NEMESCHKAL, H.L. 1991. Character coupling for taxa
discrimination: a critial appraisal of quadratic
assignement procedures (QAP). Z. zool. syst. Evolut.-
Jorsch. 29: 87-96.
176
RAUP, D.M. 1966. Geometric analysis of shell coiling:
general problem. J. Paleont. 40(5): 1178-1190.
STONE, JR. 1996. The evolution of ideas: a phylogeny
of shell models. Am. Nat. 148(5): 904-929,
TURSCH, B. 1997a. Spiral growth: The "Museum of all
Shells" revisited. J. Moll. Stud. 63: 581-588.
TURSCH, B. 1997b. Non-isometric growth and
problems of species delimitation in the genus Oliva.
Apex 12(2-3): 93-100.
VERMEU, G.J. 1993. À natural history of shells.
Princeton University Press, USA.
ROLAN
New species of Zebina
APEX 13(4): 177-179, 20 déc. 1998
À new species of Zebina (Gastropoda: Rissoidae: Rissoininae)
from Yucatän (Mexico)
E. ROLAÂN
Cânovas del Castillo 22, 36202 Vigo, Spain
KEY WORDS. Rissoininae, Zebina, new species, Yucatan, Mexico.
ABSTRACT. Zebina unamae n. sp. is described. The species was found in Yucatan, Mexico,
Caribbean Sea. It is compared with related species.
INTRODUCTION
The subfamily Rissoininae 1s represented by numerous
species in the Caribbean. They have been treated in
general books, such as CLENCH & TURNER (1950),
WARMKE & ABBOTT (1961), ABBOTT (1974), VOKES
& VOKES (1983). LEAL (1991). Diaz MERLANO &
PUY ANA HEGEDUS (1994), and in revisions such as that
Of DESJARDIN (1949), or in comparison with the
species of other areas, as in SLEURS (1989, 1993). The
nearby fauna of Brasil is mentioned in SOUZA LOPEZ et
al. (1966) and in RIOS (1985).
Numerous species have already been described in
this subfamily. Nevertheless, some new ones have been
recently added (see DE JONG & COOMANS, 1988:
FABER, 1990).
In 1994 the author was in Mexico, invited to
participate in the “Primera Reunion de Vinculaciôn
Académica sobre Tôopicos Malacolôgicos”. Some
sediment samples were collected snorkeling in front of
the Puerto Morelos Station of the Institute for Marine
Sciences and Limnology of the National Autonomous
University of Mexico (UNAM). Two shells from this
sediment were sufficiently different from any known
Caribbean species to be considered as new to science,
and are the subject of the present work.
Genus Zebina H. & A. Adams, 1854
Zebina unamae n. sp.
(Figs. 1-4)
Type material.
Holotype (Fig. 1), 1.54 x 0.94 mm. deposited in the
Museo Nacional de Ciencias Naturales of Madrid. (n°
15.05/31010), and one paratype (Fig. 2), 1.87 x 0.91
mm, in the collection of the author.
Type locality.
Puerto Morelos. Estaciôn of the UNAM, 15 km west of
Cancun, Quintana Roo, Yucatän, Mexico.
Etymology.
The specific name 1is dedicated to the UNAM, the
institution which invited us to a scientific meeting and
from which Station we collected the sediment samples
in which the shells were found.
Description.
Shell (Figs. 1-2) oval-elongate with rissoiniform
features, apex acute, with the last whorl representing
more than half of the total shell length.
Protoconch (Fig. 3) conical and smooth, of non-
planktotrophic larval type, with 172 whorls.
Teleoconch with about three to four whorls
increasing rapidly in size. The spiral sculpture 1s the
most important, consisting of very prominent cords.
These cords number five on the first whorl of the
teleoconch, decreasing to three on the second whorl; an
additional cord appears on the third whorl from the
suture below; on the last whorl there are 8-10
prominent spiral cords, with those of the base being
smaller.
Microsculpture (Fig. 4): the spiral cords are strong
but irregularly constructed, with many small holes and
lateral prolongations. Between the cords, the surface is
formed by small deep axial sulci that are irregularly
arranged, sometimes crossing and fusing to each other.
Semicircular aperture with an expanded outer lip and
deep anal sinus. Columellar lip enlarged, weakly
concave. Outer lip enlarged externally, and being a
little undulant by the end of the spiral cords. Peristome
with one undulating thread and some parallel lines on
its inner surface.
Periostracum fine, translucent and adherent.
177
APEX 13(4): 177-179, 20 déc. 1998
Discussion.
We have had some doubts about the generic
assignment Of Z. unamae n. sp. It seemed that this
species showed more indications of belonging to the
genus Stosicia, type species ÆRissoa planaxoides
Grateloup, 1838, due to its smooth conical protoconch,
strong spiral sculpture and axial microsculpture; the
external aspect of Z. unamae is even rather similar to
Stosicia annulata (Dunker, 1859), figured in SLEURS
(1996). The most important features separating Z.
unamae from the genus Sfosicia are the absence of a
broad, deep anterior channel (PONDER, 1985) and the
lack of a more or less prominent angulation at the inner
end of the anterior channel (mentioned in SLEURS,
1996), instead having a deep posterior channel, which
is shallow in Sfosicia. Furthermore, the shell of Z.
unamae has very weak, parallel threads on the inner
side of the outer lip of the aperture, typical of the
genera Zebina or Schwartziella. But in Schwartziella
the shell always has axial nbs and spiral microsculpture
(very different from the sculpture of the present shell),
and species of the genus Zebina usually have smooth,
often shining shells, only sometimes with spiral
sculpture (PONDER, 1985). In this situation more
importance was attached to the apertural features,
rather than to other characteristics, when the final
decision was made to assign the present species to the
genus Zebina.
In view of the difficulty of that decision, we prefer
not to suggest an appropriate subgenus based solely on
the shell characters.
No other species of the Caribbean region has the
spiral sculpture and microsculpture of Z. unamae n. sp.
The only species with some similarity is Rissoina
hummelincki De Jong & Coomans, 1988, but that
species has axial ribs on the first whorls of the
teleoconch and the spiral sculpture is formed by
smaller, very numerous cords.
ACKNOWLEDGEMENTS. To Jesüs Ortea and Felipe
Flores Andolais, Director of the UNAM Station of
Puerto Morelos who invited us to participate in that
meeting. To José Templado, Marta Calvo, Chefy
Âlvarez, Angel Valdés and Zoila Graciela Castellanos,
who were companions during the collecting in this
area. To my mother Margarita Mosquera, who found
the shells of this new species after many hours of
separating micromolluscs from the sediment samples.
To Colin Redfern, of Boca Raton, Florida, for his
critical revision of the manuscript and for providing
literature. To both referees, Dr. W. Sleurs and Dr. C.
Massin, for their critical comments and advice on the
present paper. To Jesüs Méndez of the CACTI of the
Vigo University for the SEM photographs.
REFERENCES
ABBOTT, R. T. 1974. American seashells (2 Ed). Van
Nostrand Reinhold Co. New York. 663 pp.
178
New species of Zebina
ROLAN
CLENCH, W. J. & R. D TURNER. 1950. The Western
Atlantic marine mollusks described by C. B. Adams.
Occasional Papers on Mollusks 1(15): 233-403.
DE JONG, K. M. & H. E. COOMANS. 1988. Marine
gastropods from Curaçao, Aruba and Bonaire. E. J.
Brill. Leiden. 261 pp.
DESJARDIN, M. 1949. Les Rissoina de l'Ile de Cuba.
Journal de Conchyliologie 89: 193-208.
DIAZ MERLANO, J. M. & M. PUYANA HEGEDUS. 1994.
Moluscos del Caribe colombiano. Un catälogo
ilustrado. Colciencias y Fundaciôn Natura, Bogotä,
290 pp.
FABER, M. J. 1990. Studies on West Indian marine
molluscs, 19. On the identity of Zurbo bryereus
Montagu, 1803, with the description of a new species
of Rissoina (Gastropoda: Prosobranchia: Rissoidae).
Basteria 54 (1-3): 115-120.
LEAL, J. H. 1991. Marine Prosobranch Gastropods
from oceanic islands off Brazil. Universal Book
Services. Oegstgeest. 419 pp.
PONDER, W. F. 1985. A review of the genera of the
Rissoidae (Mollusca: Mesogastropoda: Rissoacea).
Records of the Australian Museum, suppl. 4: 1-221.
RIOS, E. C. 1985. Seashells of Brazil. Fundaçâo Cidade
do Rio Grande, Universidade y Museo
Oceanografico. 288 pp.
SLEURS, W. J. M. 1989. A zoographical analysis of the
Rissoinine fauna of the eastern Pacific with special
reference to a comparison with the Caribbean fauna
and with a checklist of the Eastern pacific
Rissoininae Stimpson, 1865 (Mollusca: Gastropoda).
Annales Société Royale Zoologique Belgique 119(2):
155-164.
SLEURS, W. J. M. 1993. A revision on the recent
species of Rissoina (Moerchiella), R. (Apataxia), R.
(Ailinzebina) and R. (Pachyrissoina) (Gastropoda:
Rissoidae). Bulletin de l'Institut Royal des Sciences
Naturelles de Belgique (Biologie) 63: 71-135.
SLEURS, W. J. M. 1996. A revision of the recent
species of the genus Sfosicia (Gastropoda:
Rissoidae). Mededelingen van de Koninklijke
Academie voor Wetenschappen, Letteren en Schone
Kunsten van België 1: 117-158.
SOUZA LOPES, H., À. DOS SANTOS COELHO & P. de SA
CARDOSO. 1966. Contribuiçôes ao conhecimento dos
gastrépodes marinhos do Brasil. Boletim do Museo
Naciona (Zoologia)(Rio de Janeiro) 254: 1-11.
VORKES, H. E. & E. H. VOKESs. 1983. Distribution of
shallow-water marine mollusca, Yucatan Peninsula,
México. Mesoamerican Ecology Institute,
Monograph 1. Middle American Research Institute,
publ. 54. 183 pp.
WARMKE, G. L. & R. T. ABBOTT. 1961. Caribbean
seashells. Livingston Publishing Co. Wynnewood,
Pennsylvania. 348 pp.
ROLAN New species of Zebina APEX 13(4): 177-179, 20 déc. 1998
500 um
DE a
7
+
i
,
—
Figs. 1-4. Zebina unamae n. sp. Fig. 1. Holotype, MNCN. Fig. 2. Paratype, coll. E. Rolän. Fig. 3. Protoconch of the
holotype. Fig. 4. Microsculpture.
179
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Press, Chicago.
MAYR, E. 1989. Attaching names to objects. In: What the philosophy of biology is : essays for David Hull
(M. Ruse, ed.), 235-243. Klumer Academic, Dordrecht.
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